Milnor's Isotopy Invariants and Generalized Link Homotopy
tpo32三篇托福阅读TOEFL原文译文题目答案译文背景知识
tpo32三篇托福阅读TOEFL原文译文题目答案译文背景知识阅读-1 (2)原文 (2)译文 (5)题目 (7)答案 (16)背景知识 (16)阅读-2 (25)原文 (25)译文 (28)题目 (31)答案 (40)背景知识 (41)阅读-3 (49)原文 (49)译文 (53)题目 (55)答案 (63)背景知识 (64)阅读-1原文Plant Colonization①Colonization is one way in which plants can change the ecology of a site. Colonization is a process with two components: invasion and survival. The rate at which a site is colonized by plants depends on both the rate at which individual organisms (seeds, spores, immature or mature individuals) arrive at the site and their success at becoming established and surviving. Success in colonization depends to a great extent on there being a site available for colonization – a safe site where disturbance by fire or by cutting down of trees has either removed competing species or reduced levels of competition and other negative interactions to a level at which the invading species can become established. For a given rate of invasion, colonization of a moist, fertile site is likely to be much more rapid than that of a dry, infertile site because of poor survival on the latter. A fertile, plowed field is rapidly invaded by a large variety of weeds, whereas a neighboring construction site from which the soil has been compacted or removed to expose a coarse, infertile parent material may remain virtually free of vegetation for many months or even years despite receiving the same input of seeds as the plowed field.②Both the rate of invasion and the rate of extinction vary greatly among different plant species. Pioneer species - those that occur only in the earliest stages of colonization -tend to have high rates of invasion because they produce very large numbers of reproductive propagules (seeds, spores, and so on) and because they have an efficient means of dispersal (normally, wind).③If colonizers produce short-lived reproductive propagules, they must produce very large numbers unless they have an efficient means of dispersal to suitable new habitats. Many plants depend on wind for dispersal and produce abundant quantities of small, relatively short-lived seeds to compensate for the fact that wind is not always a reliable means If reaching the appropriate type of habitat. Alternative strategies have evolved in some plants, such as those that produce fewer but larger seeds that are dispersed to suitable sites by birds or small mammals or those that produce long-lived seeds. Many forest plants seem to exhibit the latter adaptation, and viable seeds of pioneer species can be found in large numbers on some forest floors. For example, as many as 1,125 viable seeds per square meter were found in a 100-year-old Douglas fir/western hemlock forest in coastal British Columbia. Nearly all the seeds that had germinated from this seed bank were from pioneer species. The rapid colonization of such sites after disturbance is undoubtedly in part a reflection of the largeseed band on the forest floor.④An adaptation that is well developed in colonizing species is a high degree of variation in germination (the beginning of a seed’s growth). Seeds of a given species exhibit a wide range of germination dates, increasing the probability that at least some of the seeds will germinate during a period of favorable environmental conditions. This is particularly important for species that colonize an environment where there is no existing vegetation to ameliorate climatic extremes and in which there may be great climatic diversity.⑤Species succession in plant communities, i.e., the temporal sequence of appearance and disappearance of species is dependent on events occurring at different stages in the life history of a species. Variation in rates of invasion and growth plays an important role in determining patterns of succession, especially secondary succession. The species that are first to colonize a site are those that produce abundant seed that is distributed successfully to new sites. Such species generally grow rapidly and quickly dominate new sites, excluding other species with lower invasion and growth rates. The first community that occupies a disturbed area therefore may be composed of specie with the highest rate of invasion, whereas the community of the subsequent stage may consist of plants with similar survival ratesbut lower invasion rates.译文植物定居①定居是植物改变一个地点生态环境的一种方式。
ESTIMATION OF SLUDGE SEDIMENTATION
ESTIMATION OF SLUDGE SEDIMENTATIONPARAMETERS FROM SINGLE BATCH SETTLING CURVESALEXIS VANDERHASSELT 1M and PETER A.VANROLLEGHEM 2*M1Laboratory of Microbial Ecology,University Gent,Coupure Links 653,B-9000Gent,Belgium and2BIOMATH Department,University Gent,Coupure Links 653,B-9000Gent,Belgium(First received 1December 1998;accepted in revised form 1April 1999)Abstract ÐIn this paper a comparison is made between two means of obtaining the parameters for the settling velocity models that are at the core of the solid ¯ux theory:(i)the traditional approach using zone settling velocity (V ZS )data obtained from a dilution experiment and (ii)a new direct parameter estimation method relying on a single batch settling curve (SBSC).For four distinct sludges,settling curves were recorded at di erent sludge concentrations (X )and the Vesilind parameters were calculated in the traditional way.The value of the resulting model was evaluated by cross-validating it on its ability to describe complete SBSC's.Provided the dynamics of the sludge blanket descent were moderate a settler model incorporating these traditional Vesilind parameters could reasonably match the experimental batch settling curves.However,when the dynamics of the sludge blanket descent were fast,the Vesilind model failed.It was tried whether other settling velocity models could result in better ®ts to the SBSC.Here,the Cho model turned out to be the most e ective one.When cross-validating the Cho model on the dilution experiment V ZS -data,however,it was found to be less performing than the Vesilind model in describing the relationship between V ZS and X .The fact that the Vesilind model is superior to the Cho model in describing such relationships,while the Cho model is better in describing complete settling curves,clearly points out that current settling models are still very empirical in nature.An important ®nding was that with all settling velocity models tested,practical identi®ability problems appeared,indicating the need for better experimental designs.Finally,the ¯ux curves associated with the SBSC-estimated Cho parameters were compared with the traditional Vesilind ¯ux curves.Although both types of ¯ux curves generally showed similar trends,the reliability of SBSC-based ¯ux curve predictions is at present insu cient to warrant replacement of the traditional estimation of settling characteristics.#1999Elsevier Science Ltd.All rights reservedKey words Ðactivated sludge,batch settling curves,¯ux curves,parameter estimation,sedimentation,settling modelsNOMENCLATUREA surface of the clari®er (m 2)V 0Vesilind maximum settling velocity (m/h)k 'Cho parameter (kg/m 2h)n Vesilind parameter (m 3/kg)n 'Cho parameter (m 3/kg)r h Takacs settling parameter associated with the hindered settling component of the settling velocity equation (m 3/kg)r p Takacs settling parameter associated with the low concentration and slowly settling component of the suspension (m 3/kg)SBSC Single Batch Settling CurveV 0'Takacs maximum theoretical settling velocity (m/h)V ZS stirred zone settling velocity (m/h)X sludge concentration (kg/m 3)X min minimum attainable sludge concentration (kg/m 3)X t threshold sludge concentration for activation of the minimal ¯ux restriction Q uunder¯ow ¯ow rate (m 3/h)INTRODUCTIONThe design and operation of secondary clari®ers is commonly based on the solid ¯ux theory (Ong,1992;Daigger,1995).The basic data required for the application of this theory can be obtained from multiple batch tests by which the stirred zone settling velocities (V ZS )over a range of sludge con-centrations (X )are measured (dilution experiment).The relationship between X and V ZS is most often characterised by the Vesilind equation [Eq.(1)]which contains two parameters:V 0and n .The key underlying assumption is that the settling velocity is only dependent on the local sludge concentration.The Vesilind parameters can then be used to con-Wat.Res.Vol.34,No.2,pp.395±406,2000#1999Elsevier Science Ltd.All rights reservedPrinted in Great Britain0043-1354/99/$-see front matter395/locate/watresPII:S0043-1354(99)00158-X*Author to whom all correspondence should be addressed.Tel.:+32-9-264-5932;fax:+32-9-223-4941;e-mail:peter.vanrolleghem@rug.ac.bestruct the ¯ux curve which is the key input of thesolid ¯ux theory.Obtaining the Vesilind parameters by a dilution experiment is unfortunately time con-suming and labour intensive and can result in scat-tered data (Ekama et al.,1997).An alternative approach consists of linking sludge volume indices as SVI,DSVI and SSVI to V 0and n through empirical functions (Daigger,1995;Ozinsky and Ekama,1995).This linking is done on the basis of extensive historical data sets on sludge volume indi-ces and Vesilind parameters.These relationships have to be used with care because the V ZS is in¯u-enced by factors not incorporated in the sludge con-centration and sludge volume indices (Ozinsky and Ekama,1995).A more structural analysis of the di erent SVI indices allowed Bye and Dold (1998)to seriously question the validity of any correlation between Vesilind parameters and sludge volume in-dices.This leaves operators and researchers at pre-sent with the choice between a labour intensive and a non-unconditional approach.Cacossa and Vaccari (1994)and Vanrolleghem et al.(1996)successfully ®tted one-dimensional settling models to single batch settling curves (SBSC).Hence,batch settling curve based parameter esti-mation appears to have the potential of being a good means of e ciently obtaining information on sludge settling characteristics for use in the solid ¯ux theory.However,each of the above studies was restricted to one speci®c kind of sludge and in neither of the mentioned papers ¯uxes calculated with the obtained parameter estimates were com-pared against ¯uxes obtained with the traditional approach using dilution experiment V ZS 's.Consequently,the work that resulted in this paper focused on the ability of SBSC-based charac-terisation of settling velocity models for use in solid ¯ux theory instead of the method based on dilution experiment V ZS -data.Attention was particularly paid to the practical identi®ability of the parameters from such SBSC data sets and to the comparison between the resulting ¯ux curves and the ¯ux curves obtained from traditional interpretation of dilution experiment V ZS -data sets.METHODSExperimental layoutThe recently developed Settlometer (Vanrolleghem et al.,1996;commercially available from Applitek nv,Deinze,Belgium)allows to automatically record complete batch sludge settling curves,and quantify the initialsettling velocity (V ZS )and the stirred sludge volume (SSV).Sludge settles in batch mode in a settling column that has a height of 70cm and a diameter of 14cm,and is equiped with a stirrer (0.3rpm).Four di erent kinds of sludge (Table 1)were brought to the laboratory where they were aerated over night before use in an experiment.In this way sludge with stable sedi-mentation characteristics was obtained (Vanderhasselt and Verstraete,1999).The storage and settling experiments were performed at room temperature.Each sludge was thickened for 1h and subsequently pumped into the Settlometer where it was allowed to settle for 35min.Next,the sludge was rehomogenised by shortly mixing with air.To set up an experiment at a lower sludge concentration,part of the sludge in the device was sub-sequently replaced by supernatant and a new settling curve was recorded.The amount of sludge to be removed at each dilution step was calculated in such a way that the obtained sludge concentrations were more or less equidi-stant.This procedure was repeated until six settling curves were recorded.Further,care was taken to ensure that at the highest sludge concentration applied still a substantial descent of the sludge blanket occurred,while at the lowest concentration,still a sharp sludge blanket/water interface could be detected.The suspended solids concentration was determined by centrifugation (NBN 366.01,1956).From the dilution experiment described above,the Vesilind par-ameters were obtained by linear least-squares regression of the log(V ZS )against the sludge concentration.Parameters obtained with this approach will be referred to as the tra-ditional Vesilind parameters.Modelling layoutIn the past,quite a number of settling velocity models have been presented in literature (Vesilind,1968;Dick and Young,1972;Takacs et al.,1991;Otterpohl and Freund,1992;Cho et al.,1993;Watts et al.,1996).On lab settler data Grijspeerdt et al.(1995)identi®ed the Takacs et al.(1991)model [Eq.(2)]to be the best.Watts et al.(1996)stated that for the prediction of sludge blanket dynamics the Takacs model [Eq.(2)]can be simpli®ed to the Vesilind model [Eq.(1)].The latter model clearly is the most frequently used (Ong,1992;Ozinsky and Ekama,1995;Bye and Dold,1998).Cho et al.(1993)also advised that the number of parameters in a model should be lim-ited to two in order to be simple enough for application in practice.In view of these re¯ections the Vesilind model was identi®ed as the most interesting to start ter in the study the two-parameter Cho settling velocity model [Eq.(3)]was applied as well.V s V 0e ÀnX1 V s V H 0 e Àr h X ÀX minÀe Àr p X ÀX min2V s kH eÀn H XXX 3Batch settling experiments were thought to be an inter-esting information source for parameter estimation as the data are the result of only the physical properties of theTable 1.``Traditional ``Vesilind parameters for the di erent sludgesSludge Concentration range (kg/m 3)V ZS range (m/h)V 0(m/h)n (m 3/kg)Correlation coe cientB 15.6±3.00.5±3.610.30.190.9693D 4.1±0.50.4±4.7 5.50.640.9712O 14±2.20.8±9.112.20.210.9677P6.6±1.00.7±3.26.90.360.9854Alexis Vanderhasselt and Peter A.Vanrolleghem396measuring device and the settling properties of the sludge.The settling device can be assumed to have constant prop-erties.There are no bulk ¯ows in the batch-wise operating device and,consequently,no associated momentums that in¯uence the movement of the sludge blanket.Additionally,model calibration problems such as the lo-cation of the in¯uent layer (Watts et al.,1996;Ekama et al.,1997)or the determination of short circuit ¯ows from the feedlayer to the sludge recycle (Dupont and Dahl,1995)are avoided.The settling velocity models were incorporated in a set-tler model discretisised into 49horizontal layers of equal volume.This discretisation can be seen as a ®nite di er-ence approximation of the underlying partial di erential equation and was introduced by Stenstrom (1975).The large number of layers was chosen to resolve the detailed behaviour of the settling process as recommended by Jeppsson and Diehl (1996).Further,this number of layers is required to obtain a resolution that is similar to the ac-curacy of the measured settling curve.The layer structure was combined with the minimal ¯ux restriction that restrains the gravitational ¯ux from a certain layer to the layer below to the minimal gravitational ¯ux of both layers (Stenstrom,1975).Vitasovic (1989)activated this restriction only when the sludge concentration reached a certain minimum concentration (X t ),i.e.3kg/m 3.Ekama et al.(1997)identify this restriction as being necessary for numerical stability of the simulation.However,there is also physical ground to this concept:a layer can not unconditionally discharge sludge to a layer in which there is already a substantial amount of sludge.To avoid that X t would in¯uence the result without being actually esti-mated,it was decided to activate the minimal ¯ux restric-tion throughout the whole concentration region (X t =0kg/m 3)for all sludges considered.This does not pose a pro-blem for the description of batch settling data because the sludge concentrations that determine the height of the sludge blanket are relatively high and the sludge concen-tration increases continuously from top to bottom.Further,for Vesilind and Cho based settler models these sludge concentrations are located in the descending part of the ¯ux curve,so the minimal ¯ux restriction will always be active regardless the fact whether X t is equal to 0or 3kg/m 3.The height of the sludge blanket was de®ned as the lo-cation of the uppermost layer with a sludge concentration higher than a certain threshold.Vitasovic (1986)and Vanrolleghem et al.(1996)used 3kg/m 3as the threshold.From a numerical simulation with di erent sludge blanket thresholds it was found that the de®nition of this threshold is not critical.Evidently,this statement only holds as long as the threshold sludge concentration is lower than the sludge concentration at the start of the ex-periment.For practical reasons this sludge blanket threshold was therefore set to 1kg/m 3,being lower than the initial sludge concentration of all modelled exper-iments.When a settling curve is recorded (Fig.1)one observes that some start-up phenomena take place before the des-cent of the blanket starts.To incorporate also these phenomena in the batch settling model,the settling vel-ocity model was slightly changed.This was done by multi-plying the settling velocity model with an on±o term.This term is 0when the simulation time (t )is smaller than the start-up time (T st )and 1when it is larger than T st .In this way the start-up phenomena are incorporated in the model without a ecting the settling part itself.Moreover,this method has the advantage that the settling velocity model is activated as soon as the blanket starts to des-cend.RESULTSBefore starting the parameter estimation,the clari®er model was evaluated from a theoretical point of view.Analysis of the concentrationpro®lesFig.1.Settling curves from the dilution experiment with the B sludge.Estimation of sludge sedimentation parameters 397showed that the evolution of the sludge blanket in the zone settling phase is determined only by the in-itial sludge concentration.Only when the settling curve starts to bend (i.e.when the settling velocity decreases)higher concentrations have a signi®cant impact on the evolution of the sludge blanket.Hence,only from that time instant onwards,the data contain information on more than one sludge concentration.Therefore,it is postulated to be an essential experimental condition that the obtained settling curve shows su cient amount of bending before reliable parameter estimation can be per-formed on a SBSC.This statement can be sup-ported from a mathematical point of view:thereisFig.2.Data and model output from the Vesilind based model for the O sludge at 11.0kg/m 3.(A)Traditional calibration on set of V ZS .(B)Optimisation result V 0=13.1m/h,n =0.219m 3/kg,T st =6.70E À02h.Alexis Vanderhasselt and Peter A.Vanrolleghem398only one additional parameter needed to describe a straight line once one point (the initial height at the starting of the sedimentation phase)of the curve is known.Settling curves at di erent sludge concentrations were recorded for each sludge.As an example the resulting settling curves for the B sludge are depicted in Fig.1.For the di erent sludges,the tra-ditional Vesilind parameters resulting from such di-lution experiment are summarised in Table 1.Looking at the parameter values one can conclude that sludges with good settling properties (B,O)as well as poor settling sludges (D,P)were included in the experimental plan.Cross-validation 1:traditional Vesilind parameters to single batch settling curveFor each experimental data set,the batch settler model was solved using the traditional Vesilind par-ameters.For T st ,a value was used that gave a mini-mal squared error (SSE)between the simulation result and the data.Provided that the dynamics in the sludge blanket were rather slow,reasonable agreement (and,thus,cross-validation)could be obtained between the data and the simulation results e.g.Fig.2(A).Still,when the zone settling phase is ending and the settling curve starts to bend [in Fig.2(A)between t =0.2and t =0.5h]asmallFig.3.Data and model output for the B sludge at 5.0kg/m 3.(A)For the Vesilind based model with traditional calibration on set of V ZS .(B)Optimisation results of the Cho based model k '=29.6kg/h m 2,n '=0.12m 3/kg,T st =1.10E À02.Estimation of sludge sedimentation parameters 399discrepancy appears between the data and the simu-lation result.For settling curves with faster dynamics (higher V ZS ),e.g.Fig.3(A),it becomes obvious that the discrepancy becomes bigger for this transition zone.Still,although the model (with Vesilind settling velocity model and parameters obtained from a dilution experiment)is not able to predict a solids pro®le that allows a smooth tran-sition between the zone settling phase and the com-paction phase,it is able to accurately describe the zone settling phase and the ®nal height of the blan-ket.Similar trends could be observed with the other sludges.It can be concluded that the model allows to adequately describe slow dynamic batch settling curves when it is calibrated with parameters from a dilution experiment.Parameter estimation from a slow dynamic single batch settling curveFor the estimation from a SBSC approach,a direction set algorithm (Brent,1973)was used to ®nd the parameters (V 0,n and T st )that would result in a minimal SSE between the observed data and the simulation results.For the data of Fig.2(A)this resulted in the ®t depicted in Fig.2(B).As can be seen for this slow dynamic SBSC,the obtained ®t is rather good.The ®t in Fig.2(B)is better than the ®t of Fig.2(A).This is as expected because the simulated curve in Fig.2(A)is not the result of a parameter estimation exercise,but the result of a cross-validation of parameters obtained from the dilution experiment.Further,the estimated Vesilind parameters (V 0=13.1;n =0.22)can be considered rather close to the ones obtained from the traditional determination (V 0=10.3;n =0.19).Although the above is important for the quality assurance of lab experiments,it is the ¯ux curves in which practitioners are interested.In order to evalu-ate this,the ¯ux curves for both parameter sets (tra-ditional vs SBSC estimation)were plotted against each other (Fig.4).As can be seen from this ®gure,both curves are rather close to each other,although in the upper concentration region di erences run up to 17%.Fig.4can also be considered as a cross-validation from the SBSC approach to the dilution approach.In the present case,the cross-validation result is reasonable.However,in other cases (data not shown)the results were less.Evaluation of practical identi®abilityWhen estimating parameters it is important to check whether the available data are informative enough to give unique values to the model's par-ameters.When the information content of a data set is too low,it is possible that di erent parameter sets can be found giving equal ®t.This practical identi®ability phenomenon surfaces when di erent estimates are found when the parameter search al-gorithm is started from di erent initial parameter guesses.In order to assess eventual practical identi-®cation problems in this application,the sum of squared errors was calculated for a large number of parameter sets in a parameter subspace.In the resulting Fig.5one observes no sharp minima but rather broad valleys.This clearly points towards identi®cation problems,i.e.it is not straightforward to ®nd the lowest sum of squared errors,leading to the best parameter set.This identi®ability problem was further con®rmed by tests that were conducted in which the search algorithm was startedfromFig.4.Flux curves for the traditional and the SBSC (X =11.0kg/m 3)estimated Vesilind parametersfor the O sludge.Alexis Vanderhasselt and Peter A.Vanrolleghem400di erent initial parameter sets,resulting in di erent optimisation results.As the SBSC approach was not successful in ®nd-ing unique parameter values,it was evaluated whether a multiple response ®tting method as used by De heyder et al.(1997)could provide a solution.In this approach,parameters are estimated using di erent settling curves obtained with di erent con-centrations of one kind of sludge.The identi®cation is then based on the fact that each settling curve contains di erent information due to the di erent initial sludge concentration and the fact that for all curves the same Vesilind parameters must hold.Note,however,that T st is dependent on V ZS (Vanderhasselt et al.,1999)and hence,it is di erent for each curve.Therefore,an optimal T st value was determined for each settling curve.Subsequently,for each settling curve the sum of squared errors was calculated in the V 0/n parameter space.The resulting error surfaces were quite similar inshapeFig.5.Simulation error as function of di erent parameter combinations for O sludge at 11.0kg/m 3.Top:n =0.219m 3/kg.Bottom:T st =0.067h.Estimation of sludge sedimentation parameters 401as the one at the bottom bining these surfaces did not result in a graph with a sharp minimum.Consequently,multiple response ®tting was not successful in solving the identi®ability pro-blem.Parameter estimation from a fast dynamic single batch settling curveFor sedimentation curves with faster dynamics it was not possible to obtain Vesilind parameters which were able to describe the curve adequately.In particular,no parameter set could be found that was able to describe the transition between the zone settling phase and the compression phase substan-tially better than the traditional Vesilind parameters [Fig.3(A)].Because this Vesilind approach was not fully suc-cessful,alternative settling velocity models as pro-posed by Takacs et al.(1991)and Watts et al.(1996)were evaluated.Neither of these models was able to describe the data substantially better than the Vesilind model.A hybrid Vesilind settling velocity model [Eq.(3)]that was proposed by Cho et al.(1993)to describe the relation between V ZS and X in dilution exper-iments,was tested subsequently.It is important to realise that special attention must be directed to the lower concentration region when this Cho model is used to calculate settling velocities:when the sludge concentration reaches zero,the settling velocity goes to in®nity,which is not physically possible.In order to prevent this,a threshold concentration below which the settling velocity is set to zero was incorporated.Arbitrarily this constant was set at 0.1kg/m 3.The precise value of this threshold was identi®ed not to be critical for this particular con-centration.One should note the correspondence between this threshold concentration and the non-settleable fraction X min in the Takacs settling vel-ocity model.Among all models tested,the Cho model gave the best ®ts to the settling curves under consider-ation,i.e.it gave the lowest SSE.The better ability of the Cho model to describe complete batch settling curves is obvious if one compares the ®t of the Vesilind model [Fig.3(A)]with the one of the Cho model [Fig.3(B)].While the Cho model pro-vided better ®ts indicating its superior structural ¯exibility,still identi®ability problems similar to the problems observed with the Vesilind model were encountered.In Fig.6the Cho ¯ux curves calculated with par-ameters obtained from di erent SBCS at di erent concentrations are depicted.Next to the Cho ¯ux curves the traditional Vesilind ¯ux curve is depicted.Considering Fig.6,it is clear that the ¯uxes obtained with the SBSC-estimated parameters should be used with precaution as there can be some di erence between the ¯ux curves obtained from parameters estimated from SBSC's at di erent sludge concentrations.For the D sludge as well as for the other sludges the highest ¯ux curves were obtained with parameters estimated from the settling curve with the highest initial sludge concen-tration.In the higher sludge concentration region the Cho ¯ux curves with parameters obtainedfromFig.6.Flux curves for D sludge,calculated with the Vesilind parameters traditionally calibrated on aset of V ZS 's and Cho parameters obtained from di erent SBSC's.Alexis Vanderhasselt and Peter A.Vanrolleghem402the SBSC approach were situated below the tra-ditional Vesilind ¯ux curve.For each sludge the ¯ux curves with the par-ameters obtained from the middle of the three selected sludge concentrations are plotted in Fig.7.It becomes clear that the ¯uxes resulting from the parameter estimation follow the same trend as observed with the Vesilind ¯ux curves obtained from dilution experiments.Sludges that have hightraditional Vesilind ¯ux curves like B and O also have high Cho-from-SBSC ¯ux curves.Sludges with low traditional ¯ux curves like D and P have corre-sponding low Cho ¯ux curves.Cross-validation 2:Cho SBSC-estimated parameters to dilution experiment V ZSIn order to further investigate the validity of the Cho model,it was evaluated whether the Chopar-Fig.7.Flux curves for the di erent sludges according to the Vesilind model traditionally calibrated on a set of V ZS 's (closed symbols)and the Cho model obtained from the middle concentration SBSC's(opensymbols).Fig.8.Natural logarithm of the V ZS in function of the P sludge concentration.Estimation of sludge sedimentation parameters 403ameters resulting from the SBSC approach could describe the X/V ZS relationships as observed in the dilution experiment(cross-validation).To this end the natural logarithm of the observed and par-ameter predicted V ZS were plotted as function of the sludge concentration,e.g.Fig.8.The Cho par-ameters resulting from the SBSC approach described the observed V ZS/X data less accurate than the Vesilind parameters from the dilution ex-periment.Also the observed correspondence was even lower for the poorly settling sludges as D and P compared to the well settling sludges.DISCUSSIONThe Vesilind parameters calculated from a di-lution experiment V ZS's allow a49layer sedimen-tation model to roughly describe complete experimental batch settling curves.However,when the settling dynamics observed in the experiment increase,the discrepancy between the simulation result and the data enlarges,especially at the tran-sition between zone settling and compression phases.On this observation di erent comments can be made:.When the sludge blanket reaches the bottom of the batch settler,compression phenomena start (or might start)to determine the descent of the sludge blanket in a direct way.In the com-pression phase sludge particles lean on each other.This is a fundamentally di erent situation than in the zone settling phase which is the only phenomenon characterised in the dilution exper-iment(Ekama et al.,1997).Here,the descent of the blanket is depending only on the equilibrium between gravitational and hydraulic friction forces..When the sludge blanket is low,it is also close to the conical bottom of the settling column.Some boundary processes related to the conical bottom and scraper could in¯uence the observed pro-cesses.A hampering of the blanket movement at the bottom could be substantiated by the®nding that in nearly all cases the¯ux curves obtained from parameter estimation on a SBSC are situ-ated below the traditional Vesilind¯ux curves at the high concentration end.The Cho model[Eq.(3)]was found to be the most e ective in®tting the observed settling curves. Grijspeerdt et al.(1995)identi®ed the Takacs model [Eq.(2)]to be the best,although they didn't include the Cho model in their comparison.Here,it has to be mentioned that the values Grijspeerdt et al. (1995)found for the Takacs r p and r h parameters are rather close to each other indicating that both exponential terms interfere with each other over quite a broad range.This is in contradiction with the mechanistic explanation Takacs et al.(1991) give to their model.Further,Grijspeerdt et al.(1995),like Takacs et al.(1991)and Watts et al. (1996),used not a settling curve but a discrete solids pro®le as experimental data for the parameter estimation.Considering the empirical nature of the settling models it would not be surprising that the optimal model structure is depending on the nature of the experiments to be described.In this respect cross-validation of parameters between the two types of data sets as performed by Cacossa and Vaccari(1994),is certainly a task for further research.Because data on concentration pro®les were lack-ing,the Cho model[Eq.(3)]was cross-validated in an alternative way:it was evaluated on its ability to describe the evolution of the V ZS as a function of the suspended solids concentration(dilution exper-iment).For the good settling sludges like the O and B sludge,the model was able to reasonably describe the observed behaviour.However,for the poorly settling sludges like P and D,the Cho model[Eq.(3)]was not able to describe the trends as well as the Vesilind model[Eq.(1)].Hence,the Vesilind model[Eq.(1)]appears to be better in describing the V ZS obtained from dilution experiments, whereas the Cho model[Eq.(3)]is better in describ-ing complete single batch settling curves.This points clearly towards the empirical nature of the used settling velocity models and on the fact that not all processes involved in the batch settling pro-cess are understood and taken into account.In the past,practical identi®cation problems as-sociated with the estimation of parameters from a single settling curve have only been addressed to a minor extent.Cacossa and Vaccari(1994)(vaguely) mentioned that the parameters estimated by their algorithm were``somewhat sensitive to the initial guesses used''.These authors stated further that reasonable parameter estimation was only possible if the sludge volume was less than half the initial volume after60min of sedimentation in their120-cm tall stirred column.However,no detailed analy-sis was reported.In the current study identi®ability problems were still encountered with curves that had a sludge volume less than a quarter of the in-itial one after30min of settling in a70-cm tall col-umn.Vanrolleghem et al.(1996)stated that no more than three parameters in a Takacs based batch settler model could be identi®ed from SBSC's recorded with the Settlometer.Despite these practi-cal identi®ability problems encountered while using SBSC's,the determination of Vesilind parameters through the more labour intensive,traditional di-lution experiment can also be troublesome as Ekama et al.(1997)report that scattered data can be obtained.From slow dynamic SBSC's one can obtain¯ux curves which are quite close to the ones obtained by a traditional dilution experiment.However,this is not always the case.For fast dynamic SBSC's the traditional Vesilind model is not able to describeAlexis Vanderhasselt and Peter A.Vanrolleghem 404。
Segue
Sloan Extension for Galactic Underpinnings and Evolution(SEGUE)Segue(v.):To proceed to what follows without pauseHeidi Newberg1,Kurt Anderson2,3,Timothy Beers4,Jon Brinkmann3,Bing Chen5,Eva Grebel6, Jim Gunn7,Hugh Harris8,Greg Hennessy9,ˇZeljko Ivezic7,Jill Knapp7,Alexei Kniazev6,Steve Levine8,Robert Lupton7,David Martinez−Delgado6,Peregrine McGehee2,10,Dave Monet8,JeffMunn8,Michael Odenkirchen6,JeffPier8,Connie Rockosi11,Regina Schulte−Ladbeck12,J.Allyn Smith10,Paula Szkody11,Alan Uomoto13,Rosie Wyse13,Brian Yanny141Rensselaer Polytechnic Institute2New Mexico State University3Apache Point Observatory4Michigan State University5ESA/Vilspa,Madrid,Spain6Max-Planck-Institut f¨u r Astronomie,Heidelberg7Princeton University8US Naval Observatory,Flagstaff9US Naval Observatory,DC10Los Alamos National Laboratory11University of Washington12University of Pittsburgh13Johns Hopkins University14Fermi National Accelerator LaboratoryI.Project SummaryA.Science GoalsWe propose an imaging and spectroscopic survey to unravel the structure,formation history, kinematics,dynamical evolution,chemical evolution,and dark matter distribution of the Milky Way.These results underpin our knowledge of the formation of the Milky Way Galaxy and of for-mation processes within the Milky Way,and will be the cornerstone of our understanding of galaxy formation processes in general.Cosmology is entering a precision era,facilitated by new work on the Cosmic Microwave Background by the Wilkinson Microwave Anisotropy Probe(WMAP)and on the distribution of galaxies by the Sloan Digital Sky Survey(SDSS)and by smaller,deeper large-telescope surveys.Galaxy formation and evolution,however,is still as data-starved as cosmology was twenty years ago.The Milky Way is the only galaxy in which we can hope to kinematically and spatially resolve the fossil record of evolution,since here the geometry is relatively unambiguous and intrinsically faint stars can be readily observed.Two key projects which focus on Galactic history and dynamics are:(1)detection of sub-structure in the Galactic halo,and(2)defining and refining our knowledge of the Milky Way’s Galactic disks.Other projects that will be necessary to realize the full potential of the key projects include understanding the relationship between SDSS photometry and the physical properties of stars,mapping the interstellar extinction in three dimensions,and studying the chemical evolution of the earliest Milky Way stars.To accomplish these goals,the existing SDSS hardware and software will be used to image (infive SDSS passbands)four thousand square degrees of the Milky Way at low Galactic latitude and to sample the stars in this new region and in the existing high-latitude SDSS survey to targetand obtain spectra of240,000stars.These spectra will allow determinations of radial velocities and chemical abundances,which will allow us to study kinematics,dynamics,and the chemical history of the Galaxy.Key Project1:Halo SubstructureSDSS data have already been used to trace the tidal stream from the Sagittarius dwarf galaxy,and to discover a second large tidal stream in the plane of the Milky Way.The structure of the Milky Way’s halo is sufficiently lumpy that it has so far defied a consistent globalfit to the smooth component of the spheroid stars.The halo may contain a(possiblyflattened)power law component,aflattened inner halo,at least two large tidal streams,a dozen dwarf galaxies and a hundred globular clusters.Some of the dwarf galaxies and globular clusters are currently being pulled apart by tidal forces.Some of the stars in the components that appear to be smooth in density may retain kinematics of the stellar associations in which they were born.Disentangling the structure of the Milky Way halo requires that individual stellar associations can be separated by population(age and metallicity),kinematics,and spatial density.The SDSS and SEGUE photometry can be used for photometric parallax(spatial density)and isochrone fitting to stellar components(into rough age and metallicity bins).Galaxy components can be separated kinematically with radial velocities.The stellar physical properties determined from spectroscopy and from imaging of open clusters serve as a check on the isochrones and further serve to illuminate the chemical composition of each component.The dynamics of stellar streams allow us tofit the Galactic potential’s shape and orientation, and constrain parameters characterizing the lumpiness of the halo dark matter.The dark matter itself could accrete with time as the stars do;knowledge of the Galactic merger history and grav-itational potential place important constraints on N-body models of galaxy formation and on the expected velocity distribution of dark matter particles.The velocity distribution of dark matter particles can affect both the energy spectrum and annual modulation of the expected signal in dark matter direct detection experiments on the Earth.Key Project2:Disk structureImaging at low Galactic latitude will allow us to study the transition from thin disk to thick disk toflattened inner halo.There is general agreement on the number and exponential form of the Milky Way stellar disks,but little agreement on the exact parameters of each.This situation is similar to the state of our knowledge of the halo several years ago,when there was general agreement that the spheroidal population of stars was well modeled by a power law,but with no agreement on theflattening parameter(measurements range from c/a=0.4−1.0).As the current generation of precise data is beginning to show,the Milky Way’s disks are not as simple as the present models suggest.The fact that the stars associated with the large tidal stream in the plane of the Milky Way were initially widely attributed to an extended thick disk of stars underlines how little we know for sure about the number and detailed form of the Milky Way’s disks.SEGUE will use spectra(physical properties and radial velocities),photometry(stellar popu-lation and spatial density),and proper motions from outside catalogs to separate and normalize the Galactic components in the solar neighborhood.Additionally,the disk components will be traced using stars as close asfive or ten degrees from the Galactic plane,using techniques similar to those forfinding halo substructure.We do not expect to be able to trace all the spiral substructure of the young thin disk,as that is best studied at longer wavelengths,but we will be able to trace the structure,kinematics,and compositions of the other disks as a function of position in the Galaxy. This latter goal will require that we understand the three dimensional structure of the Interstellar Medium(ISM),including independent measures of dust from SEGUE observations,and its effects on our stellar photometry.Formation HistoryThe identification and characterization of the Milky Way components can be utilized as an archaeological“dig”illuminating the fossil record of galaxy evolution.We will study how many mergers and of what size and time series must have occurred to make the Milky Way.We will begin to be able to pick apart the chemical and dynamical evolution of the Galaxy as a whole.We will search for rare,especially low metallicity,stars that‘freeze-in’the state of the ISM during the earliest stages of star formation in the Universe.The rare stellar objects identified in this survey will provide followup targets of high scientific interest for the world’s largest telescopes.B.Survey DataApproximately four thousand square degrees of new imaging data,at moderate to low Galac-tic latitude,and spectra of240,000Galactic stars will be acquired.The imaging footprint was designed so that no part of the sky(aboveδ=−20◦observable from the Apache Point Observa-tory)is more than10◦from either an SDSS or a SEGUE imaging stripe.In the Galactic caps, no part of the sky is more than5◦from an imaging stripe.In addition,the scans are designed to tie the photometric calibration from the SDSS north Galactic cap to the scans in the south,and to cross each other a sufficient number of times to reduce systematic uncertainties in the overall photometric calibration.The positions of the spectroscopic plates are chosen to sample the Galaxy in all directions,so that no part of the observable sky is more than about ten degrees from a spec-troscopic plate,and to target well-studied open clusters.Figure1shows the approximate layout of the SEGUE imaging stripes and spectroscopic plates on the sky.The low Galactic latitude imaging enables studies of the metal-rich Galactic thin disk,the vertical structure of the thin and the thick disks,the Galactic warp andflaring,the three dimen-sional structure of the ISM,and present star forming regions.The imaging will be taken in similar weather conditions,at the same scanning rate(which translates to the same exposure time),and with the same instruments andfilters as the SDSS.Each stripe is2.5◦wide and requires two interleaved scans with the SDSS imaging camera,on separate nights,to complete.The imag-ing includes twelve constant Galactic longitude stripes which go through the Galactic plane and typically extend thirty-five degrees on either side(dashed green lines in Figure1).These stripes are separated by about20◦of Galactic longitude,varying somewhat to pass through known open clusters,SIRTF legacyfields,and patches of low extinction near the Galactic plane.The constant Galactic longitude stripes connect and overlap SDSS imaging of the Galactic caps to facilitate the photometric calibration of both old and new data.In addition,three new SDSS stripes(72,79,and90,shown as solid green lines in Figure1) will be imaged in the South Galactic Cap.Only three stripes were imaged in the South Galactic Cap during the SDSS survey,and the additional stripes are needed to sample that part of the sky about every ten degrees.One stripe(red line in Figure1)follows the Sagittarius dwarf tidal stream across the northern sky.Two long(half-filled)strips(magenta lines in Figure1)cross the remaining SEGUE stripes,and will be used to cross-calibrate the stellar photometry to a level of at least2%(systematic+rms),and will trace low latitude structures,including the newly discovered tidal stream in the Galactic plane.The SDSS camera must scan along great circles,so all of these stripes describe great circles on the Celestial Sphere.The spectroscopic observations include1200spectra in each of200individual sky directions. The plate positions were chosen to sample the sky in all observable directions,and spectra will be selected to sample stars from one to100kpc from the Sun and from as many Galactic substructures as possible.Additional observations target regions of high interest such as open clusters,star formation regions,and known tidal streams in the halo.Each plate position is3◦in diameter.We will design two640-fiber plates in each plate position,with a total of about1200stellar targets-60-300306090120150180210240270300-90-60-300306090-90-60-30030609010230130l b s90s86s79s72s9s16s27s37s4418h22h2h 6h 6h 17.4h 2718.0h 618.6h 837.9h 498.6h 1621.1h -022.8h 25 1.3h 32 3.6h 17 5.0h -11Figure 1.Low Latitude Imaging and Spectroscopy Plan.The SFD (1998)reddening map is shown in Galactic coordinates;note the center is shifted to (l,b )=(120◦,0◦).Green,red,magenta (purple)lines indicate new SEGUE scans to be obtained.SEGUE Imaging at l =110degrees from −30◦<b <30◦has already been obtained as of Nov 2003(SDSS runs 4134,4135,4144,4152).The red line along the great circle with (node,incl)=(32◦,35◦)follows the Sagittarius dwarf tidal stream.Magenta lines indicate half-filled “strips”in portions of the sky at low Galactic latitude,and cross the constant longitude stripes for better calibration;the great circles arcs are defined by (node,incl)=(259.9◦,43.6◦),(311.0◦,66.7◦).The total SEGUE imaging area is about 4000sq.degrees,of which 200sq.degrees has already been obtained.Black lines indicate existing or planned SDSS regular imaging.Black dotted reference lines are at b =0◦and declination (DEC)=−20◦(no SEGUE imaging or spectroscopy occurs at a DEC of <−20◦from Apache Point Observatory in the Northern hemisphere).Black long dashed lines mark constant Right Ascensions (RAs)of (18,22,2,and 6)bels above the top of the figure indicate RA,DEC start and end for a vertical SEGUE imaging stripe.Open black circles indicate positions of known Sagittarius dwarf tidal stream stars.Filled black circles indicate positions of known Monoceros stream stars.Open black diamonds indicate positions of known Galactic open clusters.The blue circles indicate individual 3-degree diameter positions of Galactic structure plate pairs (bright plate:45min exposure,plus faint plate:90min exposure),168blue plate pairs.Yellow circles indicate positions of ‘special plates,’landing on a known open cluster,the Sag.dwarf stream,or the Monoceros Ring structure.29blue plate pairs.Total:197plate pairs and about 240,000stellar spectra with resolving power R ∼2000,and 3<S/N <100for objects with 20.3>g >14.5.plus calibration objects.One plate will have the SDSS standard 45minute exposure time,and the other will be exposed for twice as long,allowing us to reach stellar targets as faint as g ∼20.3.Spectroscopic targets will include halo giants,metal-poor dwarfs,G disk and halo dwarfs,white dwarfs,and a large variety of rare stars.At low latitudes,targets within star-forming regions will be selected.II.Scientific CaseOur understanding of Galaxy evolution has advanced considerably since the monolithic col-lapse model of Eggen,Lynden-Bell and Sandage(1962)was adopted as the standard.Most experts now believe that the Galaxy was built up through a series of mergers(Searle&Zinn1978),though there is no agreement on the number and size of the merger events.These current models of galaxy formation stem from cold dark matter(CDM)simulations that show the outer halos accret-ing over billions of years(Steinmetz&Navarro2002),and from the increasing number of examples of moving groups and tidal disruption discovered in the halo of our galaxy(Majewski et al.2003; Newberg et al.2002;Odenkirchen et al.2001a;Irwin&Helmi et al.1999;Ibata,Gilmore,& Irwin1995;Grillmair et al.1995;Irwin&Hatzidimitriou1995),M31(Ferguson et al.2002),and other external galaxies(Shang et al.1998,Zucker et al.2004).It is possible that the hierarchical merging process is most important in the dark matter dominated galactic halos,while disks might form from the(angular momentum conserving)collapse of the gas within the stellar spheroid.However,Λcold dark matter simulations suggest merging may significantly affect the formation of disks as well(Abadi et al.2003).Evidence of mergers is currently most apparent in the outer halo where signatures of satellite accretion persist for many gigayears(Johnston,Spergel,&Hernquist1995).Dwarf galaxies and globular clusters are among the outer halo objects which are merging at the current epoch.It is also possible that there exist some lumps of dark matter in the outer halo that have not yet merged,and which do not contain stars(e.g.,Bullock,Kravtsov,&Weinberg2001).These latter structures could be evident by their perturbation of tidal tails and warping of disk structures.Our own Milky Way is the only galaxy that we can presently study at sufficiently high spatial and kinematical resolution,and at sufficient depth,to address many of the open questions of galaxy formation and small-scale structure evolution in sub-halos.Our goal is to obtain the spectroscopic and photometric data required to unravel the structure,the formation history,the kinematic and dynamical evolution,the chemical evolution,and the distribution of the dark matter within and around the Milky Way.We propose two key projects,which contribute to our knowledge of the Galactic mass assem-bly and disk formation models.These projects are:(1)detection of substructure in the Galactic halo,and(2)defining the structures of the Galactic disks.These are really two parts of the one key project to define the major components of the Milky Way galaxy,but are listed separately since they may require different data sets and analysis methods.One may think of this proposed project as providing a large homogeneous input data set to a21st century model of the Galaxy–one which involves not only accurate multi-color photometry such as has gone into earlier models (Bahcall&Soneira1984),but large amounts of kinematic velocity and proper motion data which can be used to complete the dynamical and evolutionary picture.This technique is most similar to that used to construct the Besancon model of the Galaxy(Robin et al.2003),but with more input data.Detection of substructure in the Galactic halo requires photometry and radial velocities in as many directions as possible.The large tidal streams that have already been discovered are more than4kpc across,setting the scale over which we must sample the sky tofind all large tidal structures.Characterizing the Galactic disks requires data collection primarily at low latitudes, and within a few kpc of the Galactic plane.The goal is to separate disk components by their stellar content,and then measure the global properties.These projects will separate and describe components using radial velocities,proper motions, chemical composition,photometric parallax,and isochronefitting to photometry.In some sense, our survey is concentrated on the“big picture”of our galaxy.We will identify and constrain all of the largest components,paving the way for future inquiries which willfindfiner substructure,S52-32-20.4S341+57-22.5S297+63-20.S6+41-20S223+20-19.4S200-24-19.8S167-54-21.5(RA,DEC)[l,b](0,0)[96,-60](15,0)[128,-63](30,0)[157,-58](45,0)[177,-49](60,0)[190,-37](75,0)[199,-25](90,0)[207,-11](105,0)[214,2](120,0)[221,15](135,0)[229,28](150,0)[239,41](165,0)[254,52](180,0)[276,60](195,0)[308,63](210,0)[337,58](225,0)[357,49](240,0)[10,37](255,0)[19,25](270,0)[27,11](285,0)[34,-2](300,0)[41,-15](315,0)[49,-28](330,0)[59,-41](345,0)[74,-52]15171921g *23171921231517192123151719212305101520253035404550Figure 2.Polar histogram of color-selected turnoffstars on the celestial equator .This data was obtained with the Sloan Digital Sky Survey.The radial distance gives the apparent dereddened g ∗magnitude.The angular position gives the RA.Galactic coordinates are labeled for reference.The shading of each cell indicates the relative number of counts of stars in each (r,θ)=(g ∗,α)bin.A typical absolute magnitude for stars with these colors is M g ∗=4.2.The feature at α=60◦is an artifact from imperfect reddening correction of a large dust cloud at this position.The streak at α=229◦represents stars in the globular cluster Palomar 5.The boldface labels indicate our names for identified structures of halo stars.S341+57-22.5at g ∗∼22.5,and S167-54-21.5at g ∗∼21.5are cross sections of the Sagittarius dwarf galaxy tidal stream.more accurately determine the chemical evolution,and measure proper motions for a large fraction of the stars in the Milky Way.A calibrated catalog of images,spectra,and associated derived quantities,will be the primary product of this survey.These will be generated in nearly real time,to be used for rapid follow-up work or as input targets to space-based or large aperture telescopes.The need for this global picture of our galaxy is well illustrated by the results of Newberg et al.(2000;Figure 2).In this figure,there are seven marked concentrations of stars.Concentrations S341+57-22.5and S167-54-21.5have been identified as cross sections through the tidal stream ofthe Sagittarius dwarf spheroidal galaxy,which is currently in the process of tidal disruption.The overdensities S223+20-19.4and S200-24-19.8are thought to be pieces of another tidal disrupting stream in the plane of the Milky Way galaxy.The concentrations in this region near the Galactic plane,at15th and17th magnitude near the anticenter are not named,but also are not understood in any global picture of the Milky Way.The overdensity S297+63-20.is thought to be another tidal stream,possibly associated with the Sagittarius dwarf galaxy,though this has not been confirmed and remains controversial.The concentrations S6+41-20and S52-32-20.4are thought to be portions of the stellar spheroid,though their density distributions do notfit standard spheroidal models within the errors of our density measurements.One sees in Figure2a strong argument for a global view of the whole Milky Way,including low Galactic latitudes,since one cannot identify substructure without understanding the major Galactic components in which that substructure is embedded,and properly accounting for inter-stellar extinction.Many of the scientific analyses that we anticipate will be based on these data have counterparts in the much smaller-scale efforts of individuals or groups,which,unavoidably, dilute their impact by acquiring data in a piecemeal and non-uniform fashion.A uniform survey creates a synergy which allows more global questions to be addressed and leaves behind a legacy data set against which future data sets will be compared.A better understanding of the Milky Way’s structure and evolution is already a“cornerstone”project in ESA’s science planning,through the GAIA satellite mission,and plays an important role in the definition of the science goals for NASA’s SIM and TPF missions,which are“key elements in NASA’s Origins Program.”We demonstrate here how a deep imaging and spectroscopic optical survey will complement as well as lay the groundwork for these ambitious satellite projects. Furthermore,SEGUE willfill a unique and vital niche complementing ongoing and planned large ground based Galactic structure programs such as RAVE and K.A.O.S.A.Characterization of the Component Stellar Structures in the Milky WayThe fossil record of galaxy evolution(star formation and mass assembly)is written in the chemical,kinematic,and spatial distribution of Galactic stars.The main recognized components of the Galaxy are the thin disk,the thick disk,the bulge,and the stellar spheroid.Recently many groups of astronomers have identified examples of Galactic structure that either requires additional components or an increase in the complexity of the traditional components.Kinematic studies show the existence of moving groups and coherent streams(numerous studies),and a group of stars(Gilmore,Wyse,&Norris2002)that may be part of the merger that puffed up the thick disk.Overdensities of stars over the Galactic bar(Parker,Humphreys,&Larsen2002)have been found in photometry.Also,a new metal-weak thick disk component has been proposed(see Norris 1994,Beers et al.2002and references therein).Clearly,even the basic stellar components of the Milky Way are not yet understood in depth. The complex substructure now being identified has undoubtedly biased the previous limited studies of thick disk structure,contributing to our present imprecise knowledge of the thick disk;study of many lines of sight over much of the sky will be necessary to unravel the substructure and obtain a more complete picture.This survey would specifically target the thick disk/halo boundary and substructure.The structures would be studied in stellar density from statistical photometric parallaxes,and in kine-matics through statistics of the radial velocities/metallicities in each component.We use the term ”statistical photometric parallax”to describe the method of using photometry to determine dis-tance(photometric parallax)in cases where the number of stars is large,so that statistics can be used to estimate the underlying spatial structure of the group.We will have a unique opportunity to study the stellar Metallicity Distribution Function(MDF),especially in the region where the thick disk and spheroid populations overlap.Figure3demonstrates preliminary results from SDSS-4-3-2-101[Fe/H]100010000100000Z D i s t a n c e (k p c )Figure 3.The distance and metallicity distribution of EDR stars.The distance distribu-tion of ∼4000stars from the SDSS Early Data Release (EDR)as a function of metallicity [Fe /H].One can clearly discriminate the presence of thick-disk stars with metallicities in the range -1.0<[Fe/H]<0and locations within a few kpc of the Sun,from the halo objects at large distances that extend to much lower metallicities.EDR spectra.Flaring and Warping of the Galactic PlaneThe disks of many galaxies both flare and warp in their outer regions.Flaring is attributed to an increasing ratio of spherically distributed dark mass to disk mass with increasing distance from the center of the galaxy,and provides one of the few available methods of measuring the three-dimensional distribution of dark matter within a galaxy.The origin of Galactic warps is still something of a mystery.Tidal interactions with satellites and neighbors is an obvious cause;for example,the warp in the Galaxy is often attributed to the tidal influence of the Magellanic Clouds (e.g.,Weinberg and Nikolaev 2001;Garcia-Ruiz et al.2002)or of the Sagittarius dwarf spheroidal galaxy (Bailin 2003).However,not all warped galaxies appear to have (presently detected)neigh-bors.The warp andflare in the outer Galactic disk has been studied primarily using(radio)obser-vations of neutral hydrogen.The depth and color sensitivity of SDSS will allow the3-D structure of the northern warp in the Galaxy to be traced to distances beyond the entire known extent of the warp(20kpc,Binney1992),using photometrically identified giants,carbon stars(especially when combined with2MASS data)and red clump stars(e.g.,Margon et al.2002;Helmi et al.2003). The Structure of the Thick DiskHawley et al.(2002)show that early M dwarfs can be traced to distances of up to1kpc above the Galactic plane,well into the domain where the thick disk population dominates that of the thin disk.Since the stars of the thick disk are more metal-deficient(typically by at least0.3-0.5 dex)than the thin disk stars,their colors,especially g−r,diverge from those of the metal-rich disk stars.One can therefore,at least in a statistical sense,separate the two populations.Afirst look at the vertical structure of the thick disk from SDSS data has been carried out by Chen et al.(2001). Star counts in the thin and thick disks can be used to determine the initial mass function,and in particular the counts of lower-metallicity stars must be understood(see the recent discussion by Zheng et al.2001and Chabrier2003).The current SDSS imaging data provide star counts at high Galactic latitude only;under-standing and disentangling the vertical structure of both the thick and thin disks requires data covering the whole range of Galactic latitude at many longitudes.The Structure of the Galactic HaloSDSS data have already demonstrated the presence of very large structures in the Galactic halo(Yanny et al.2000;Ivezic et al.2000;Odenkirchen et al.2001a,b;Newberg et al.2002; Rave et al.2003;Yanny et al.2003).These structures include extra-tidal features around globular clusters and vast comoving stellar streams from accreted dwarf galaxies.These structures,and similar more tenuous analogues,may cover a significant fraction of the sky.Imaging more sky allows such structures to be traced to larger angular sizes,and allows structures which do not completelyfill a great circle on the sky to be detected.Furthermore,high-metallicity globular clusters tend to be found at lower Galactic latitudes;there may be streamers and tails of different color(metallicity)at lower latitudes.Indeed,recently Frinchaboy et al.2004showed that many low latitude open and globular star clusters are likely to be associated with a single large tidal stream in the Galactic plane.There is increasing evidence that even“globally recognized”structures,such as“the halo,”change dramatically with increasing Galactocentric distance.An inner,“flattened”halo compo-nent,for example,has been indicated by many recent studies(Lemon et al.2003,Chiba&Beers 2000).Preston,Shectman&Beers(1991)have used the mean colors of blue horizontal-branch stars to indicate a possible decrease in the ages of stars with increasing Galactocentric distance. Sirko et al.(2004)use an analysis of the spectra of high-latitude A stars to demonstrate a change in the velocity ellipsoid of the halo with distance from the Galactic center.Both of these represent key results for understanding the formation of the Milky Way,which could be readily refined and extended using our proposed survey effort.Complexity of Galactic spheroid populations in other galaxies is traced by their globular cluster systems(at least in early-type galaxies with populous cluster systems that can be studied). Typically the globular cluster systems are bimodal,with bluer(lower metallicity)clusters mak-ing up a more extended and dynamically hotter system,and redder(higher metallicity)clusters comprising a system more centrally concentrated and sometimes rotating(e.g.many papers in IAU Symposium207,ed.Geisler,Grebel&Minniti2002).Typically thefield-star populations have color and spatial distributions suggesting properties more similar to(and perhaps having。
辣椒细胞质雄性不育系及其保持系的AFLP分析英文
A FL P A n a lys is o f C y top la sm ic M a le 2s te rile L in e a n d Its M a i n ta i n e r L i n e o f P ep p e rLUO Xia ng 2d o n g1,2,DA I L ia n g 2fa n g1,2,CH EN J i n 2fen g23,WAN G S h u 2b i n31.C o ll ege o f Ho rti cu l tu re,N an ji ng Ag ri cu lt u ra lU n i vers it y,N an ji ng 210095;2.Co ll eg e o fL i fe Sc i ence s,J i an gxi No r m al Un i ve rs i ty,N ancha ng 330022;3.Vege tab l e R es ea rch I n s ti tu t e ,J i an gs u Academ y o f Ag ricu lt u ra l Sc i ence s ,N an j ing 210014Ab s t ra ct [O bject i ve]The a i m o f t h is study wa s t o ana l yze the cyt o p l a s m i c m a l e s teri le l i n e 21A and its m ainta i ne r li n e 21B o f p ep pe rs by AF L P,an d l ay t he fo unda ti on fo r further s tudies o n m o l ecu l a r m echan is m o f t he cyt op l a sm i c m ale s te ril ity i n p epp ers .[M ethod ]C yt op l a sm i c m a l e s teril it y (CMS)li ne 21A and its m a i n t a i ne r li ne 21B w ere a nalyzed by AFLP t o o bta i n the s p ecific am p li fied fra gm en ts o f cyt op lasm i c m a l e s teri le li ne 21A,wh i le the sp ecifi c amp li fi ed fragm en ts w ere recove red o r sequ enced,and ana l yzed by BLAS Tn and TBLASTX i n GenB a nk o f NCB I .[Result ]A sp eci fi c fragm en t AA /CAG 169w as ob tained from cytop l am i c m al e ste rili ty (C M S )li ne 21A.Pa rti a l DNA s equen ce of AA /CAG 169w as t he se ries repe ated s equen ce i n AFLP m a rker seq uence related t o the sex and grow t h tra i t s o f Pen aeu s m ono do n.The re su l ts o f BLASTX and TBLASTX showed t ha t the tran s l a ti o n p r o duct s o f t h is sp eci fi c fragm ent had a hi g he r si m il a rity to So l anum dem is sum pu t a ti ve gag 2po l p ro tei n ,So l anum dem i s sum p u t a ti ve retro tra nspo so n p r o te i n and So l a num d em is sum p uta ti ve re tro tran spo so n gag p ro t e i n re s p ec ti vel y.[C on clu si on]The functi o n o f this spec i fic fragem ent AA /CAG 169is mo re li ke l y t o be cl o s el y rel a ted t o re tr o transp o son,wh i ch i nd i cate s tha t the sp eci a l fragm en t is po ss i b l y re l a t e d t o the s te ril ity trait .Key w o rds Pep pe r;cy t op l a s m i c m ale 2s t e ri lti y (C M S);AFLP ana l y sisR e ce i ve d:J a nua ry 11,2010 Accep t e d:Fe b rua ry 23,2010Suppo rted by Na ti o nal 863Prog ram i n t h e 11th Five 2ye a r P l an ni n g Pe ri o d (2006AA1001082322);Na ti ona l Na tural Science Fo unda ti o n (36;3);S T y j f x O ff f (G 5);Y G F f x N U y (3)32jf @j A s one of vege tab l e c rop s with the l a rge s t cu lti va ti on a re ai n C hi na ,p epp e r (Ca ps i cum a nnuu m L .)is a wo rl d w ide ki nd of vege ta bl e s and p roc e ssing condi m e nts .Acco rdi ng t o the sta tis ti c s,a nnua l c ultiva ti o n a rea of peppe rs i n C hi na ha s re ac he d abo ve 1400tho usa nd h m 2,whi ch m a ke s gre a t con 2tri buti o n to annua l a ve ra ge ba lance supply of vege ta bl e s inChina [1].W ith obvi o us adva nta ge s in p re 2m a turity,hi gh yi e ld a nd fru itw e i ght,peppe r hybri d s a re pop ul a r w it h m any fa r m 2e rs .It is expe nsive t o bree d hyb ri d see ds by a rti fic i a l em a scula ti on a nd po lli na ti on .I t is l a bou r 2sa vi ng t o ob t a i n hy 2bri d se ed s F 1by cytop l a sm i c m a le ste ril e li ne of peppe rs,which ca n e nsure the p urity o f se e ds and p ro t e c t the ri ghts of bree de rs,a nd is a lso the m o st e ffi c ient wa y t o re duce se e dco st a t p re se nt [2]o r the m a j o r direc ti o n fo r curre nt p epp e r bree ding .Thus,be side s e nha ncem en t o f the i de nti f i ca ti o n a nd b re ed i ng of c yt opl a s m i c m a l e ste ril e li ne o f peppe rs,stud 2i e s shoul d be ca rried out a t di ffe ren t l eve ls from b i o l ogy,which w ill p r o vi de a theo re tica l ba sis f o r furthe r effi c i e nt o r w i de u tili za ti on .R ul e s of inhe ritance in m o r p hol ogy,cyt o l ogy,POD isozy m e a nd m a l e 2ste ril e cha ra c t e rs ha ve be en a lrea dy dis 2cu ssed t ho roughl y i n studi e s o n t he m e cha nis m of cyt op l a sm i cm a le ste ril e li ne of peppe rs [3-4].Howe ve r,the re is little report on the m o l e cul a r m e chan is m of ste rility i n pe ppe rs,e spe c i a ll y se que nce fra g m e nts re la ted to cyt op l a s m ic m a l e s te ril ity of peppe rs .The re fo re ,ge nom ic DNA fr om c ytop l a s m i c m a le ste ril e l ine 21A a nd its m a i n ta i ne r l ine 21B o f peppe rwa s com 2pa ra ti ve l y ana lyze d by AFLP t o ob ta i n t he spe cifi c am p l ifi e d fragm en ts re l a ted to CM S i n this s tudy,wh i ch l a id a fo unda 2ti on fo r fu rthe r studi e s on mo l e cula r m e cha nis m of the cyt o 2pla sm ic m a l e ste rilit y in p epp e rs,a nd pro vide d a refe re nce fo r be tte r utili za ti o n of cyt op l a s m ic m a le ste rility i n peppe rs .M a te ria ls a n d M e th o d sMa te ria l sThe cyt op l a s m ic m a l e ste ril e line 21A w ith comp l e t e l y s ta 2ble ste rility tra its a nd its m a inta i ne r l ine 21B of peppe rs we re from Ve ge table Re se a rch I ns titute ,J i a ng su Aca dem y of Agri 2c ult u ra l Sc i e nce s .S ee dli ngs of 21A a nd 21B we re bred on the coo l ing bed i n e a rl y J a nua ry,w hil e i n l a te Ma rch,100p l a nts of 21A a nd 21B we re re spe c tive l y p l a nte d with conve nti ona l fie l d m a na gem e nt i n p l a s ti c gre e nho use s of Ve ge tab l e R e 2sea rch I ns titute ,J i a ng su Aca dem y of Ag ri cultura l S c i e nc e s .Me thod sA FL P R e fe rence d by Vo s [5]a nd LOU Q un 2feng [6],re stri c 2ti on fragm e nts a nd do uble 2stra nd DNA we re j o i nte d t oge the r w ith T 4li ga se through EcoR I and M se I doubl e di ge sti o n,w hil e re stri c ti o n e ndo nuc l e a se a nd T 4l iga se w e re a ll p urc ha se d from S ha ngha i S angon B i o l ogi c a l Engi ne e ri ng Te chno l o gy &S e rvi ce s Co .,Ltd .5μl d i ge sti o n p roduc t wa s used a s the tem p l a te f o r p re 2am p li fi ca ti on,w hil e the p ri m e rs of p re 2am p li fi 2c a ti on we re E +1a nd M +1.5μl p roduct of p re 2am p l ifi ca ti on wa s a na l yzed by 1%aga ro se ge l e l e c tropho re sis to e sti m a te the e ffi c iency of p re 2amp li fi c a ti on .Acco rding t o the te sti ng fo r p r o duc ts o f p re 2am p li fi c a ti on,the pro duc ts w e re dil ute d 10-30fo l d,a nd t he n u se d a s the tem pla te fo r se l e cti ve amp li fi c a 2ti on .F i na ll y,p roducts o f se lec tive amp li fica ti o n w e re sepa ra 2ted by 6%dena tured po l ya c ryl am i de ge l e lec tr opho re sis a nd dye d by s i lve r sta i ni ng .Re c yc ling o r c lon ing a nd se que nc e a na lys is of sp ec i f ic b and s D i ffe ren ti a l ba nds of cyt op l a sm i c m a le ste ril e li ne a nd its m a i nta ine r li ne we re re co ve re d a nd cut off a ccura te l y by the z f μ22S f 5f f 5μ2f 2Ag ri cu lt u ra l Science &Techno l o gy,2010,11(2):69-71C op yri gh t κ2010,I n f o r m a ti on Ins titute of HAAS.A l l right s re s erved.Agricultura lB i o techno l o gy0801200900781cience and e ch no l o g P ro ec t rom J i a ng i P ro vi nc i a l i ce o Educa ti on J J 0819o un g r ow thu nd o J i ang i o r m o l n i ve rs it 298.C o rre spo n di n g au tho r .E m ail :chen n au.e ste rili e d sca l pe l t o p ut i nto c en tri uga l tube s w ith 200l re d is till e d w a t e r .am p l e s w e re p l a ce d i n t o bo ili ng w a te r o r 1m i n a nd then cen tri uge d o r 10m i n w ith 12000rpm.l supe rna tan tw a s use d a s the temp l a t e o r com bina ti on t o the correspo nd i ng pri m e rs aga i n w it h the sam e am plifi ca ti o n .Afte r the m olec ul a r we i ght o f amp li fied p r o duc ts wa s ve ri fi e d,and t he n li ga ted t o pGE M Tea sy ve ctor .Com pe te nt E .co li JM 109wa s transfe rre d by the li ga ted p roduc ts fo r b l ue 2wh i te se l e c ti on.S ing l e 2co l ony p l a sm i d wa s e xtrac te d f o r e ndo nuc l e a se re a c 2ti on,and the n se nt to S ha ngha i I nvi troge n B i ote chno l o gy Co .,Ltd fo r DNA se que nc ing .The who l e sequence of j o i nts we re rem oved a fte r c l oning a nd se que nc i ng,a nd then subm i tted t o NCB I da ta ba se fo r BLAS T .R e s u lts a n d A n a lys isA FL P sp ec if i c f ra gm e n t s c ree n ing of g enom e from cyto 2p lasm ic m a le s te rile line 21A a nd its m a in ta in e r li n e 21BGenom ic DNA from cyt op l a sm ic m a l e ste ril e li ne 21A a nd its m a i nta ine r line 21B w e re am plifi e d w it h 32pa i rs o f AFLP pri m e r com bi na ti ons,a nd the re sults showe d tha t bands in AFLP m ap we re ri c h w ith 60bands i n ea ch l a ne.O nl y amp li 2fi e d p roduc ts from o ne pa i r of pri m e r had di ffe rence be t w e e n cyt op l a s m i c m a l e s te ri le li ne and its m a inta i ne r li ne i n 32p a irs of AFLP p ri m e rs,wh i ch indica te d tha t the re wa s l ittl e diffe r 2e nc e in ge nom i c DNA be t we en c yt op l a s m i c m a l e ste ril e li ne 21A a nd its m a i n ta i ne r li ne 21B.Genom ic DNA from cyt op l a sm ic m a l e ste ril e li ne 21A a nd its m a inta i ne r l ine 21B we re am p li fi e d a ga i n w ith p ri m e r com bi 2na ti on EcoR 2AA /M se I 2CAG scree ned by poly mo rphic pri m e r pa irs .The re wa s one spe c i fi c band w ith 200bp in cyt op l a sm i c m a le ste ril e li ne 21A (F ig .1),w hil e no ba nd wa s f o und i n its m a inta i ne r li ne 21B.Aft e r the repe a te d te st,the fra g m e ntswe re demo nstra te d to be sta bl e ,re li a bl e and a sp ec i fic fra g 2m e nt re la ted t o m a l e 2ste rile c ha rac ters.A:Am p li fi cati o n p rodu cts o f p ri m e r EAA /M C AG,and the arr ow show s the d i ffe ren ti a l fragm e nts;B :S i ng l e c l o ne i d entifi ca ti on o f d i ffere ntial fragm e nts .F i g.1 Am p l i fi ca ti o n of AFLP d i ffe ren ti a l fragm e nts o f ge nom i cDNA from s t e ril e l ine 21A and its m ai n tai n er l ine 21BRe c yc ling o r c lon ing a nd se qu enc i n g of A FL P d i ffe re n ti a l f ra gm e ntsF rom F i g .1-B ,re com bi nan tp la s m id PCR p roduc ts we re i den ti c a lw ith the si z e of ta rge t fra g m ents,which show ed tha t ta rge t fra g m e nts ha d a lre ady be en succ e ssfu l ly T 2A cl one d .Afte r ve cto r se que nce a nd AFLP p ri m e r com bi na ti o n AA /CAG sequence we re rem ove d,the se que nc i ng sequence w a s 169bp i n ful l length (F i g .2).Thu s,the spe c i fi c f ragm e nt i n c ytop la s m i c m a l e ste ril e li ne 21A wa s m a rked a s AA /CAG 169t o ana l yze co nvenientl y.The und erli ned pa rt s are AA /CAG se quence s of AFLP p ri m e r com b i na ti o n ,wh il e the do uble 2u nde rli ned pa rts are AF LP j o i nt s .F i g.2 Sp eci fi c fragm en t sequ ences o f s teri le li ne 21ABLA S Tn a na lys is of AA /CAG 169s equ en ce in sp ec ific f ra g 2m en tsAcco rdi ng t o BL AS T re sults,pa rti a l sequence s of AA /CAG 169(l oc a ted in 122-158bp ,AGA ATTGGT A C G 2CAGTC T AT G ATG AGT CC T G AGT A AC ,w ith 37bp i n tota l)had a h i ghe r si m il a ri ty t o seve ra l sequence s o f t he re l a te d AFLP m a rke r se que nc e s with the sex and grow th tra its in P ena eu s mo nodon (AFE549M 27.1).Furthe r ana l ys is re 2ve a led tha t 37bp se que nce s we re four se ri e s re pea ted se 2que nce s of t he re l a te d AFLP m a rke r se que nc e s w it h the se x a nd grow th tra its in Pe nae us m ono don (F i g .3),so the spe 2c i fi c fra g m ent AA /CAG 169w a s cl o se l y re l a te d t o its ste rilit y .BLA S TX a na lys is of AA /CAG 169s equ en ce in sp ec ific f ra g 2m en tsA A /CAG 169se que nc e wa s subm itt e d to GeneB a nk,a nd a na l yze d by BLASTX o f NCB I a nd p rote in da ta ba se s fo r ho 2mo l ogy sea rching .The re sults sugge sted tha t am ino a c i d se 2que nce deduce d by this fra g m ent had a hi ghe r s i m ila rit y t o pa rtia l se que nce s of So l a nu m dem i s sum p uta tive ga g 2po lp ro 2t e i n a nd So l a num dem issu m puta ti ve re tr o tra nsposon p r o te in re sp ec ti ve ly (F i g .4).T BL AS TX a na l ysis i ndica te d tha t am i no a c i d se que nce deduce d by this fragm en t ha d a lso a hi ghe r y q f S 2,S S ,2F i g.3 A li gnm en t of t he pa rti al seque nce s of AA /C AG 169andthe re l a t e d AFLP m arke r se quence s with the s ex andgrow th t raits i n Penaeu s m o no done nco de d by gag (g roup 2sp ec ifi c a ntige n )ge ne i n the struc 2tura l ce n t e r of p lan t re tr o tra nspo so n,a nd the n be cam e se v 2e ra l kind s of l e ukocyte p ro te i n s a fte r pos t 2tre a t m e nt .Gag 2po l p r o te i n wa s e ncoded by po l 2ga g i n com m on,a nd the n be 2,f 2z y T f ,f f f f 2G 6y y 07Ag ri cultura l Sc i ence &Tech no l o gy Vo l .11,No.2,2010si m il a rit t o pa rti a l se uence s o o l anum dem is sum pu t a ti ve gag po l p rote i n o lanu m dem issum puta ti ve re tro tra nspo so n pro te i n a nd o l a num dem issu m puta ti ve re tr o tra nsposon ga g pro te i n .Howe ve r gag p ro te i n wa s a m ulti p r o te in p re curso rc am e re ve rse tra nsc ri p t a se inte gra se a nd pro te a se a te r en m e di ge sti on.he re ore the unc ti o n o this spe c i ic ra ge m e n t AA /CA 19wa s m o re like l to be c l o se l re la te d t o re tro tra nspo son .F ig.4 B LASTX a nalys i s o f t he w ho le sequ ence o f C M S 21A 169D is c u s s io n sM e chanism of m a l e s te ril ity ge ne s ha s bee n w i de l y stud 2i ed a t hom e a nd a broa d,a nd it is ge ne ra lly conside red tha t the o ccu rrence of plant CMS i s c l ose ly re l a te d t o the struc ture a nd exp re s si o n of m i tocho ndria l ge nom e ,whil e the frequent re com bi na ti on o f intra 2m o l e cul a r a nd i nte r 2mo l e cula r m i tochon 2dri a l genom e is m o l e cul a r ba sis f o r the occu rre nce o f C M S.F rom a l a rge amo un t of m it o chondri a l DNA fragm e nts i n the i denti fi e d CM S ,the y a re a ll gene ra ll y ch i m e ri c ge ne s from pa rtia l sequence s o f se ve ra l i den ti fi e d ge ne s afte r m ulti 2re 2com bi na ti on a nd the un i de nti fi e d re a di ng fram e (urf),such a ska l e nap us [7],m a ize [8]a nd ri ce [9].The t o ta l ge nom i c DNA e xtrac te d by C T AB i s cons titute d by nuc lea r DNA,m t D NA a nd cpDNA,so the spec ifi c fra g m ent AA /CAG 169of ste ril e li ne 21A po ssibl y com e s fr om nuc l e a r DNA,and is a lso p rop e rl y from m t DNA o r cpDNA.The ope n re ad i ng fram e of the sp ec i fic fra g m e nt A A /CAG 169se que nce is found w ith the on l ine t oo l of O RF F i nde r in NCB I .Com bi ne d w it h BLAS T X and T BLASTX a na l ysis,the op en rea di ng fram e of the sp ec i fic fra g m e nt AA /CAG 169se que nce is pri m a ril y i denti fi e d to be -1encodi ng f ram e fo r 53am i no a c i ds.W he re a s,the true i de nti f i ca ti o n of t h i s re a di ng fram e is still unknown,a nd its re l a ti onshi p w i th ste ril e chi m e ri c ge ne s nee ds furthe r studi e s .BL AS Tn in this study re ve a ls tha t the spe c i fi c fra g m e nt AA /CAG 169is li ke l y t o be c l o se l y re l a te d t o ste rili ty .Howe ve r,BLAS T X and T BL AS T X a ll show tha t AA /CAG 169sequence is po ssibl y re l a te d t o re tro tra nspo son,which i ndica te s tha t c yt op l a s m i c m a l e ste rility o f peppe rs m a y be c l o se l y re l a te d t o re tr o tra nspo son .The si m il a r re sults a rea lso repo rted i n cyt op l a s m ic m a le ste rilit y of ca rro ts [10],whi c hl a ys a founda ti on fo r furthe r s t udie s on m o l ec ul a r m e cha nis mof cyt opla s m ic m a l e ste rilit y in peppe rs .R e fe re n c e s[1]F ANG R (方荣),CHEN XJ (陈学军),M I A O NS (缪南生),et a l.Adva nce s i n gene ti c r e sou rces and mo l ecular breed i ng o f pepp er (Cap s i cum spp.)(辣椒(Cap s i cum spp.)遗传资源与分子育种进展)[J ].Acta Ag ri cult u rae J i angxi (江西农业学报),2004,16(3):55-61.[2]WANG SB (王述彬),ZHAO HL (赵华仑),L I U J B (刘金兵),et a l.He t e r o s is util i za ti o n o f C MS li ne s i n ho t (swee t )p eppe rs and t e ch 2ni q ues fo r i ts hybri d seed p roducti o n (辣(甜)椒胞质雄性不育系杂种优势利用及其制种技术)[J ].J i ang su J ou rnal o f Ag ri cultura l Sci 2ences (江苏农业学报),2002,18(3):143-146.[3]DA ILF (戴亮芳),LUO XD (罗向东),WA N G S B (王述彬),et a l.I so zym e s anal y ses o f cyt op l a s m i c m a l e s teri l e (C M S )l i ne i n pep 2p er (C ap s i cum annuum L.)(辣椒细胞质雄性不育系的3种同工酶分析)[J ].Ac t a Bo tan i ca B o reali -Occi den t a li a Si nica (西北植物学报),2007,27(9):1772-1776.[4]LUO X D ,DAI LF ,WANG SB ,et a l .Ma l e g am ete deve l o pm entand ea rl y t ape t a l degene rati on in cyt op l a s m i c m a l e ste ril e pepp er (Cap s i cum annuum L.)i n ve st i g ated by m e i o ti c,ana t o m i ca l and ultras tructu r a l ana l yse s[J ].Pl an t B reed i ng,2006,125:395-399.[5]V O S P,H O GER R ,BLEEKER M.AF LP:a n ew techni qu e f o rDNA fi ngerp ri n ti ng [J ].Nucl e i c Ac i ds R es,1995,23:4407-4414.[6]LOU QF (娄群峰),CHEN J F (陈劲枫),JAHN M,et al .I den ti fica 2ti o n of AF LP and SC AR mo l ecu l ar m a rke rs li nked t o gyno eci ou s l o 2ci in Cucum i s s ati vus L.(黄瓜全雌性基因连锁的AF LP 和SCAR 分子标记)[J ].Acta Ho rti cult u rae Si n i ca (园艺学报),2005,32(2):256-261.[7]Y ANG J H,ZH AN G M F,Y U JQ.M it ochond ri a l n ad2gene is co 2tran scri p ted w i th CMS 2as soci a ted o rf B ge ne i n cyt op l a s m i c m al e 2s teri l e s tem m us t a rd (B ra s si ca j uncea)[J ].Mo l ecu l a rB i o l o gy Re 2po rts,2009,36:345-351.[8]GALLAGHER LJ ,B ETZ SK,CHASE CD.M it ochond ri a l R NA edi 2ti n g trunca t e s a ch i m eri c op en reading fram e as sociated w i th S m ale 2s t e ril it y i n m a i ze[J ].C urr Gene t,2002,42:179-184.[9]ZHANG H,L I SQ ,YIP,e t al .A Hon gli an C MS li ne of rice dis p l aysabe rran t F 0of F 0F 12ATPas e [J ].P l ant C e l l R epo rts,2007,26:1065-1071.[10]NAK AJ I MA Y,Y A MA MO TO T,MUR ANAK A T,et a l .Gene ti c var 2i ati o n o f p etal o i d m al e 2ste ri l e cyt op l a s m o f ca rro ts reveal ed by se 2que nce 2t agged s i tes (STS s )[J ].Theo r App l Gene t ,1999,99:837-843.R es p o n s i b le ed ito r:C H EN Xiu 2c h en R es p o n s i b le t ra n s la to r:L I J in g 2w e i R es p o n s i b le p ro o fread er:W U Xiao 2y an辣椒细胞质雄性不育系及其保持系的AFL P 分析(摘要)罗向东1,2,戴亮芳1,2,陈劲枫23,王述彬3 (1.南京农业大学园艺学院,江苏南京210095;2.江西师范大学生命科学学院,江西南昌330022;3.江苏省农业科学院蔬菜研究所,江苏南京210014)[目的]对辣椒细胞质雄性不育系及其保持系进行AFLP 分析,为进一步深入研究辣椒细胞质雄性不育的分子机制奠定基础。
Tc-99m-MIBI serve as a predictor of chemotherapy response in malignant lymphomas compared with P-gP
British Journal of Haematology ,2001,113,369±374Technetium-99m-sestamethoxyisobutylisonitrile scan as a predictor of chemotherapy response in malignant lymphomas compared with P-glycoprotein expression,multidrug resistance-related protein expression and other prognosis factorsChia-Hung Kao,1Shih-Chuan Tsai,2Jhi-Joung Wang,3Yung-Jen Ho,4Shung-Tai Ho 5and Sheng-Ping Changlai 61Department of Nuclear Medicine,Taichung Veterans General Hospital,Taichung,2Department of Nuclear Medicine,Show-Chwan Memorial Hospital,Chunghua,3Department of Medical Research,Chi-Mei Medical Centre,Tainan,4Department of Radiology,Jen-Ai Hospital,Taichung,5School of Medicine,National Defence Medical Centre,Taipei,and 6Department of Nuclear Medicine,Chung-Shan Medical and Dental Hospital,Taichung,TaiwanReceived 13November 2000;accepted for publication 14January 2001Summary .The purpose of the present study was to predict the response of malignant lymphomas (MLs)to chemotherapy usingtechnetium-99m methoxyisobutylisonitrile (Tc-MIBI)scan and to compare it with the predictive ability of P-glycoprotein (P-gp)expression,multidrug resistance-related protein (MRP)expression and other prognosis factors.Twenty-five ML patients were enrolled in this study prior to initiation of chemotherapy .Images were obtained 10min after intravenous injection of Tc-MIBI,interpreted visually and the tumour-to-background (T/B)ratios calculated.Immunohistochemical analyses were performed on sections of the biopsy specimens to determine P-gp and MRP expression.Chemotherapy response was evaluated in the first 1±2years after completion of chemotherapy .The mean T/B ratio of the 15patients with agood response (3´3^0´6)was significantly higher than that of the 10patients with a poor response (1´2^0´1).All 15patients with a good chemotherapy response had positive Tc-MIBI scan results and negative P-gp and MRPexpression.All 10patients with a poor response had negative Tc-MIBI scan results and either positive P-gp or MRP expression.Other prognosis factors showed no significant difference in the incidence of good and poor responses.Tc-MIBI scan results represent P-gp or MRP expression more accurately than other prognosis factors and predict the chemotherapy response in ML patients.Keywords:malignant lymphoma,technetium-99m methoxy-isobutylisonitrile,chemotherapy response,P-glycoprotein expression,multidrugresistance-related protein expression.Chemotherapy is the primary therapeutic modality for many malignant lymphomas (MLs)including all non-Hodgkin's lymphoma (NHL)cases and many cases of Hodgkin's disease (HD)(Barr et al ,1997;Neal &Hoskin,1997;Wilson &Chabner,1998).As resistance to chemotherapeutic agents is a major cause of treatment failure,the goal of chemotherapy for ML is to avoid possible resistance and achieve the highest response.The mechanism of tumour uptake of technetium-99m methoxyisobutylisonitrile (Tc-MIBI)may involve bindingto the cytosol of the tumour cell (Hassan et al ,1989).The cationic charge and lipophilicity of Tc-MIBI,mitochondrial and plasma membrane potentials of tumour cells,and cellular mitochondrial content can all play a significant role in tumour uptake of this agent (Chiu et al ,1990),or the uptake may be caused by indirect phenomena such as increased tumour blood flow and capillary permeability .Tc-MIBI scans have been used to successfully predict the chemotherapy response of MLs (Kapucu et al ,1997;Shih et al ,1998).However,the previous studies have not compared the relationship between Tc-MIBI scan resultsq 2001Blackwell Science Ltd369Correspondence:Chia-HungKao,MD,Department of Nuclear Medicine,TaichungVeterans General Hospital,160Taichung Harbour Road,Section 3,Taichung407,Taiwan.E-mail:kaoch@ .twand P-glycoprotein(P-gp)or multidrug resistance-related protein(MRP)expression in predictingthe chemotherapy response of MLs.Therefore,the aim of this study was to compare Tc-MIBI scan results,immunohistochemical ana-lyses of P-gp and MRP expression,and other prognosis factors as predictors of chemotherapy response in ML patients.PATIENTS AND METHODSPatients.Twenty-five patients(13men,12women;age range25±65years;mean age:46´2^12´3years)with ML(11with HD and14with NHL)were included in the study and underwent Tc-MIBI scans prior to chemotherapy (Table I).The classification of ML followed the Lukes and Butler and updated Kiel systems(Jaffe,1998).After Tc-MIBI scans,the11HD patients received chemotherapy regimens with nitrogen mustard(mechlorethamine),vincristine, procarbazine and prednisone(MOPP),alternatingwith doxorubicin,bleomycin,vinblastine and dacarbazine (ABVD);the14NHL patients received chemotherapy regimens with cyclophosphamide,doxorubicin,vincristine and prednisone(CHOP)protocols(Barr et al,1997;Neal& Hoskin,1997;Wilson&Chabner,1998).Technetium-99m methoxyisobutylisonitrile scan.The ima-ging procedure began30min after oral intake of500mg of perchlorate to prevent any abnormal uptake of free Tc-99m pertechnetate.A commercial MIBI preparation(max. 5´56GBq in approximately1±3ml)was obtained from The Du Pont Merck Pharmaceutical Company(Cardiolite, Billerica,MA,USA).The labellingand quality control procedures were carried out accordingto the manufac-turer's bellingefficiencies were all.95%. Each patient was place in a supine position on the imaging table with the chest strapped to prevent motion. Because of physiological Tc-MIBI accumulation in abdom-inal and pelvic organs,visualization of MLs located in abdominal and pelvic regions is unreliable.In this study, images of supradiaphragmatic MLs were obtained10min after intravenous injection of740MBq Tc-MIBI in the anterior and posterior projection.The equipment consisted of a large field-of-view gamma camera fitted with a low-energy,high-resolution collimator.A single20%energy window was set at140keV and500K counts were obtained for each static image.Tumour-to-background(T/ B)ratios were calculated as the mean counts over the region of interest(ROI)of the tumour outlined in the largest lesion4the mean counts over the ROI ofTable I.Detailed data of patients in this study.Case Tc-MIBI scan results Immunohistochemical stainingAge BChemotherapyresponsenumber Sex T/B ratio Visual interpretation P-gp expression MRP expression(years)Type Stage symptoms results1Female1´0Negative Positive Negative33HD I Yes Poor2Female1´0Negative Negative Positive53NHL III No Poor3Female1´1Negative Positive Negative40HD II Yes Poor4Male1´1Negative Positive Negative51NHL III No Poor5Male1´1Negative Negative Positive62HD IV No Poor6Male1´2Negative Positive Negative35NHL IV No Poor7Male1´2Negative Negative Positive55HD IV Yes Poor8Female1´3Negative Positive Negative27NHL III Yes Poor9Male1´3Negative Negative Positive65NHL III Yes Poor10Male1´4Negative Positive Negative43NHL II Yes Poor11Female2´4Positive Negative Negative43HD II Yes Good12Male2´7Positive Negative Negative56HD IV No Good13Male2´8Positive Negative Negative37NHL III No Good14Male2´9Positive Negative Negative55HD I Yes Good15Male2´9Positive Negative Negative62HD IV No Good16Male3´0Positive Negative Negative61NHL III No Good17Female3´2Positive Negative Negative31NHL III No Good18Male3´2Positive Negative Negative47HD IV No Good19Female3´3Positive Negative Negative60HD III Yes Good20Male3´3Positive Negative Negative25NHL II Yes Good21Female3´6Positive Negative Negative30HD II Yes Good22Female3´6Positive Negative Negative58NHL IV No Good23Male4´0Positive Negative Negative35HD IV No Good24Female4´1Positive Negative Negative50NHL III No Good25Male4´5Positive Negative Negative42NHL II Yes GoodHD,Hodgkin's disease;NHL,non-Hodgkin's lymphoma;P-gp,P-glycoprotein;MRP,multidrug resistance-related protein;Tc-MIBI, technetium-99m methoxyisobutylisonitrile;T/B,tumour-to-background.370 C.-H.Kao et alq2001Blackwell Science Ltd,British Journal of Haematology113:369±374background,defined as the contralateral normal side for the neck and axilia lesions or normal soft tissue of the thorax for mediastinal lesions.Tc-MIBI uptake in the lesions $axillary soft tissue background,based on the visual interpretation of at least two experienced nuclear medicine physicians,was considered a positive Tc-MIBI scan result (Figs 1and 2).Immunohistochemical staining.Formalin-fixed paraffin sections (5m m)were deparaffinized in an oven at 508C for 40min,then hydrated with varyingconcentrations of ethanol±water dilutions.For MRP immunohistochemical staining,antigen retrieval was performed by treatment in citrate buffer in a 700W microwave oven for 5min.Endogenous peroxidase was blocked by 3%hydrogen peroxide for 15min,followed by 5min in phosphate-buffered saline (PBS).The sections were incubated over-night in a moist chamber at 48C with primary antibody MRP QCRL-1(10m g/ml,Signet Laboratories,Dedham,MA,USA)at 1:100concentration.For P-gp immunohis-tochemical staining,endogenous peroxidase was blocked by 3%hydrogen peroxide for 15min.Antigen retrieval was performed by treatment with enzyme digestion in 0´1%trypsin in PBS for 5min at room temperature and inhibited with 10%skimmed milk in PBS for 5min.The sections were incubated for 2h in a moist chamber at 378C with primary antibody JSB-1(50m g/ml,Boehringer Mannheim Biochemica,Germany)at 1:50concentration.After three 5min washes in PBS,detection of the primary antibody was performed with a link antibody accordingto the manufacturer's instructions (DAKO LSAB_2System,Peroxidase,Dako Corporation,Carpinteria,CA,USA)(Niehans et al ,1992;Marie,1995;Yamaguchi et al ,1995;Kostakoglu et al ,1998;Webb et al ,1998).P-gp and MRP expressions were interpreted by an experienced pathologist blind to clinical outcome as follows:negati-ve less than 10%,positive 10%or more stained tumour cells (Figs 3and 4).Chemotherapy response evaluation.In this study ,the chemotherapy response of each patient was evaluated for the first 1±2years after completion of treatment using clinical and radiological methods such as plain chest X-ray ,chest computerized tomography (CT)or magnetic resonance imaging (MRI),as well as head and neck CT or MRI,accordingto the followingscale:(1)Complete response no evidence of disease,(2)Partial respon-se at least 50%decrease in the sum of the products of the maximum perpendicular diameters of all measurable lesions,no evidence of progression in any lesion and no new lesions,(3)No response less than 25%increase in the sum of the products of the maximum perpendicular diameters of all measurable lesions,no evidence of progression in any lesion and no new lesions,and (4)Progressive disease at least 25%increase in the sum of the products of the maximum perpendicular diameters of all measurable lesions and/or the appearance of new lesions.We defined complete and partial responses as good response,while no response and progressive disease were defined as poor response.Statistical analyses.The T/B ratio was expressed as meanT a b l e I I .D i s t r i b u t i o n s o f T c -M I B I s c a n r e s u l t s ,P -g p e x p r e s s i o n ,M R P e x p r e s s i o n ,a g e ,t u m o u r t y p e ,t u m o u r s t a g e a n d B s y m p t o m s r e l a t e d t o c h e m o t h e r a p y r e s p o n s e r e s u l t s .I m m u n o h i s t o c h e m i c a l s t a i n i n gC h e m o t h e r a p y r e s p o n s e T c -M I B I s c a n r e s u l t s P -g p e x p r e s s i o nM R P e x p r e s s i o nA g e T y p e S t a g eB s y m p t o m sr e s u l t s P o s N e g P -v a l u eP o s N e g P -v a l u e P o s N e g P -v a l u e#40y e a r s .40y e a r s P -v a l u e H D N H L P -v a l u e I ±I I I I I ±I V P -v a l u e Y e s N o P -v a l u e G o o d1500150155107851069P o o r 010,0´0164,0´0146,0´01460´73460´74370´86640´33q 2001Blackwell Science Ltd,British Journal of Haematology 113:369±374Tc-MIBI Predicts Chemotherapy Response in HD and NHL371^standard deviation (SD).A Mann±Whitney U -test was used to evaluate the difference in T/B ratios between patients with a good versus a poor response.The difference in incidence of good and poor response was evaluated for eight possible prognosis factors:positive versus negative Tc-MIBI scan results,positive versus negative P-gp expression,positive versus negative MRP expression,HD versus NHL,stage I±II versus stage III±IV ,age .40years versus #40years,and with B symptoms versus without B symptoms (night sweats,fever .388C for three consecutive days and unexplained weight loss of .10%body weight)(Barr et al ,1997;Neal &Hoskin,1997).A Chi-square test was used to determine if the frequency of good and poor response was the same for each pair.If the P -value was ,0´05,the difference was considered significant.RESULTSDetailed patient data are shown in Table I.The mean T/B ratio of the 15patients with a good response (3´3^0´6)was significantly (P ,0´01)higher than that of the 10patients with a poor response (1´2^0´1).All 15(100%)patients with a good response had positive Tc-MIBI scan results and negative P-gp and MRP expression.All 10(100%)patients with a poor response had negativeTc-MIBI scan results,six (60%)of whom had positive P-gp expression while the other four (40%)had positive MRP expression.Tc-MIBI scan results,P-gp expression and MRP expression all showed significant differences in the rate of good and poor responses.However,no significant difference in the incidences of good and poor responses was found for lymphoma type,stage,age or B symptoms (Table II).DISCUSSIONOur review of previous literature found only one paper that reported that 17ML children with positive Tc-MIBI scan results and a higher mean T/B ratio had a better response to chemotherapy than seven ML children with negative Tc-MIBI scan results and a lower mean T/B ratio (Kapucu et al ,1997).Our results support their findings.However,their study did not examine the relationship between other prognosis factors,P-gp or MRP expression,and chemo-therapy response.The mechanism of chemotherapy resistance in ML is thought to involve expression of P-gp and MRP (Niehans et al ,1992;Yamaguchi et al ,1995;Zhan et al ,1997;Webb et al ,1998).The retention of Tc-MIBI in tumour cells depends on P-gp and MRP expression and they function as ATP-dependent efflux pumps for manychemotherapyFig 1.Case no.25had a goodchemotherapy response result.(A)Tc-MIBI scan reveals significant tracer uptake in the right neck and the result is positive (T/B ratio 4´5)(arrow).(B)Neck computerized tomography shows a mass in the same area(arrow).Fig 2.Case no.4had a poorchemotherapy response result.(A)Tc-MIBI scan reveals no definitely abnormal tracer MIBI uptake in the chest and neck and the result is negative (T/B ratio 1´1).(B)Gallium-67citrate scan shows multiple abnormal tracer uptake in the chest and neck (arrows).372 C.-H.Kao et alq 2001Blackwell Science Ltd,British Journal of Haematology 113:369±374agents (Hendrikse et al ,1998,1999;Vergote et al ,1998;Sun et al ,2000).Therefore,in this study we used the Tc-MIBI scan to predict the response of MLs to chemother-apy .We found that positive Tc-MIBI scan results accurately predicted all good chemotherapy results,which were also related to negative P-gp and MRP expression.Moreover,negative Tc-MIBI scan results accurately predicted poor chemotherapy results in all patients with positive P-gp or MRP expression (Table I).In our previous studies,only early chest images performed 10min after intravenous injection of Tc-MIBI proved to be accurate enough to predict chemotherapy response in lung and breast cancer (Kao et al ,1998,2000;Vergote et al ,1998).Therefore,in this study ,we did not consider it necessary to perform delayed chest imaging to calculate the tumour washout rate or retention index of Tc-MIBI to predict the chemotherapy response.mRNA expression is not fully corrected with P-gp or MRP expression in the tumour cell membrane,Tc-MIBI tumour uptake is directly based onthe P-gp or MRP expression in the tumour cell membrane,and it was impossible to extract mRNA from the formalin-fixed paraffin sections of biopsy specimens (Wang et al ,1997;Dexter et al ,1998;Yokogami et al ,1998).Therefore,we directly detected P-gp or MRP expression using immunostainingto correct with Tc-MIBI tumour uptake (T/B ratio)in our study .Based on our findings,we conclude that Tc-MIBI scan results can represent P-gp and MRP expression for predict-ingthe chemotherapy response in ML patients.However,further studies includinglarg er case numbers and patients who have relapsed followingchemotherapy are necessary to confirm our findings.ACKNOWLEDGMENTSThis work was supported in part by grants from Taichung Veterans General Hospital (TCVGH-896708D)andNationalFig 3.Immunohistochemistry performed on sections of malignant lymphomaspecimens from two different groups reveals (A)negative and (B)positive P-gp expression (Â500).Fig 4.Immunohistochemistry performed on sections of malignant lymphomaspecimens from two different groups reveals (A)negative and (B)positive MRP expression (Â1000).q 2001Blackwell Science Ltd,British Journal of Haematology 113:369±374Tc-MIBI Predicts Chemotherapy Response in HD and NHL 373Science Council(NSC89±2314-B-075A-015,89±2320-B-075A-001,88±2314-B-075A-006),Taiwan.REFERENCESBarr,L.,Cowan,R.&Nicolson,M.(1997)Haematological Malignancies.In:Oncology(ed.by L.Barr,R.Cowan&M. 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合并危险因素的高钾血症诊治进展
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The impact of cosmic neutrinos on the gravitational-wave background
A.
Metric perturbations in the Poisson gauge
In this paper we consistently account for the presence of cosmic neutrinos to analyze their impact on the evolution of the second-order gravitational-wave background. At linear order it has been shown that there is a damping effect due to the anisotropic stress of free-streaming neutrinos that strongly affects the primordial gravitationalwave background on those wavelengths which enter the horizon during the radiation dominated epoch (at the level of 30%) [15, 16, 17, 18, 19, 20, 21] (see also Ref. [22]). At second order, along with the analogous damping effect, we find that free-streaming neutrinos are an important source for the second-order gravitational-wave background during the radiation-dominated epoch. We find completely new source terms, arising because of the fact that neutrinos give a relevant contribution to the total energy density during this epoch and they behave as ultrarelativistic collisionless particles after their decoupling: their high velocity dispersion acts as an extra source for the second-order gravitational waves. To compute such a contribution we evaluate the second-order tensor part of the neutrinos’ anisotropic stress tensor, that has been neglected so far. This is achieved by computing and solving the Boltzmann equation for neutrinos. Approximating the neutrino contribution as a perfect fluid of relativistic particles during the radiation era leads to a serious underestimate of their role. Let us stress that the new contribution is at least of the same order of magnitude as that computed by adopting a fluid treatment in the source of the scalar-induced gravitational waves. Moreover it has a clear physical interpretation. It arises in the Boltzmann equation from a “lensing” effect of the neutrinos as they travel through the inhomogeneities of the gravitational potential.
翻译——精选推荐
细胞分子生物学文章第十卷(2005),711-719 pl2005.7.15寄出200510.6收到脂质体:一项先进制造技术的概述新西兰,北帕默斯顿,专用邮袋11222,梅西大学Riddet中心,M.REZA MOZAFARI摘要:近几十年来,脂质体作为生物膜的理想模型,也是药物、诊断、疫苗、营养物和其他生物活性剂的有效载体,引起了广泛关注。
在不同背景下研究者们对脂质体学领域的文献报道广泛地不断地增加,这表明这一领域引人入胜。
自从大约40年前脂质体被介绍到科学界,许多技术和方法在或大或小的脂质体制造规模上得到发展。
这篇文章将在大体上提供脂质体制备方法优缺点的概览,特别强调在我们实验室开发的加热法,作为一种脂质囊泡快速生产的模式技术。
关键词:载体系统,加热法,脂质囊泡,脂质体学,制造技术引言脂质体科学技术是一个正在飞速发展的科学,举几个例子,它用于诸如药物递送,化妆品,生物膜的结构和功能,探索生命起源等领域。
这是由于脂质体有一些有利的特性,例如,它不仅能包含水溶性药物也能包含脂溶性药物,在体内识别特定靶向位点,在流动性、大小、电荷、层数的方面具有多样性。
脂质体作为生物膜模型的应用限于在实验室中研究,它们在生物活性剂的包载和递送的成功应用不仅取决于脂质体载体可以达到预期目的的优越性的示范,还取决于技术和经济可行性的规划。
对于递送应用,脂质体配方应该具有高包封率,窄粒度分布,持久稳定性和理想的释放特性(根据预期的应用)。
这些要求制备方法有产生脂质体的可能性,且脂质体可采用多种成分分子,例如:脂质/磷脂可提高脂质体稳定性。
除了上述特性,对于蛋白质、核酸之类敏感的分子/化合物的递送,脂质体也应该能保护复合制剂,防止其退化。
尽管在脂质体上进行了大量的研究开发工作,但只有少数脂质体产品已被批准为人类使用至今。
这也许有许多原因:一些脂质体配方的毒性,分子和化合物在脂质体中的低包封,脂质体载体的不稳定性,脂质载体的不稳定性,特别是大尺度的脂质体生产成本高。
2018考研英语长难句解析(二)
The environmentalists, inevitably, respond to such critics. The true enemies of science, argues Paul Ehrlich of Stanford University, a pioneer of environmental studies, are those who question the evidence supporting global warming, the depletion of the ozone layer and other consequences of industrial growth. 词汇突破: 1.environmentalist 环保主义者 2.Inevitably 不可避免的 3.depletion of the ozone layer 臭氧层破坏(这属于是环保的标配词汇) 4.consequences of industrial growth ⼯业增长的后果 5.critics 批评 6.pioneer 先驱 7. question 质疑 8. argue 认为 结构分析: 共两句,第⼀句⽐较简单就不解析了。
The environmentalists, inevitably, respond to such critics. 环保主义者们不可避免地对这样的批评做出了回应。
第⼆句:主⼲识别: argues Paul Ehrlich of Stanford University, = Paul Ehrlich of Stanford University argues 主谓充当的插⼊语: The true enemies of science are those 其他成分:a pioneer of environmental studies, 同位语 who question the evidence supporting global warming, the depletion of the ozone layer and other consequences of industrial growth. 定语从句 翻译点拨: 切分之后调整语序(按中⽂习惯) 1.Paul Ehrlich of Stanford University (is)a pioneer of environmental studies, 2.Paul Ehrlich of Stanford University argues 斯坦福⼤学的Paul Ehrlich是环境研究⽅⾯的先驱, (这样诡异的不知道读⾳的⼈名在考场上⼀般就直接抄英⽂)他认为, 或者翻译为: 环境研究⽅⾯的先驱之⼀,斯坦福⼤学的Paul Ehrlich认为, 3. The true enemies of science are those 科学真正的敌⼈是这样⼀些⼈。
非等位基因
非等位基因概述非等位基因是指同一基因座上的不同等位基因。
等位基因是指在某个给定的基因座上,可以存在多种不同的变体。
每个个体继承了一对等位基因,一对等位基因可能会导致不同的表型表达。
非等位基因的存在使得遗传学研究更加复杂,因为不同的等位基因会对个体的表型产生不同的影响。
背景在生物学中,基因座是指染色体上一个特定的位置,该位置上的基因决定了某个特征的表达方式。
每个基因座上可以有多种不同的等位基因。
等位基因是指在某个特定基因座上的不同基因变体。
每个个体都会继承一对等位基因,通过这对等位基因的不同组合,决定了个体的表型。
然而,并非所有基因座上的等位基因都具有相同的表现型。
非等位基因的影响非等位基因的存在导致不同等位基因会对个体表型产生不同的影响。
有些非等位基因会表现出显性效应,也就是说,当个体继承了一个突变的等位基因时,即使同时继承了一个正常的等位基因,但显性效应会使得突变的等位基因的表型表达得到体现。
相反,有些非等位基因会表现出隐性效应,当个体继承了两个突变的等位基因时,才会表现出突变的表型。
除了显性和隐性效应之外,非等位基因还可能发生两种其他类型的表型效应。
一种是共显效应,当个体继承了两个不同的突变等位基因时,在表型表达上会表现出一种新的特征,这个特征并不是单个突变等位基因所能导致的。
另一种是部分显性效应,当个体继承了两个不同的突变等位基因时,表型表达将介于两个单独突变等位基因的表型之间。
重组和非等位基因重组是指两个不同的染色体交换部分基因序列的过程。
在重组的过程中,非等位基因可能会发生改变,导致新的等位基因组合形成。
这一过程使得非等位基因的表型效应更加复杂,因为新的等位基因可能将不同基因座的效应组合起来。
非等位基因的重要性非等位基因对生物的适应性和多样性起着重要作用。
通过对等位基因的各种组合的研究,人们可以更好地理解基因与表型之间的关系,并揭示遗传变异对物种适应环境的重要性。
总结非等位基因是指同一基因座上的不同等位基因。
辽宁高校联盟英语学位考试真题
The primary goal of the environmental protection project is to _____.A. reduce pollution levelsB. increase industrial productionC. promote tourismD. expand urban areasWhich of the following is NOT a characteristic of a successful entrepreneur?A. Strong leadership skillsB. Risk-averse natureC. Innovative mindsetD. Ability to adapt to changeThe study of _____ focuses on the relationship between language and society.A. sociologyB. psychologyC. linguisticsD. anthropologyIn the field of computer science, _____ refers to the process of organizing and storing data.A. programmingB. data structureC. software developmentD. algorithm designThe theory of plate tectonics explains the movement and interaction of _____.A. atmospheric layersB. Earth's magnetic fieldsC. large sections of Earth's crustD. ocean currentsC(正确答案)Which of the following is a key principle in sustainable development?A. Maximizing immediate profitsB. Meeting present needs without compromising future generationsC. Rapid industrializationD. Unlimited resource extractionThe term "novel" originated from the Italian word "novella," which means _____.A. a short storyB. a historical accountC. a poemD. a philosophical treatiseIn economics, the concept of "supply" refers to the _____.A. total amount of goods and services producedB. desire to purchase goods and servicesC. government regulation of marketsD. study of consumer behaviorWhich of the following is a fundamental branch of physics that deals with the nature and properties of light?A. MechanicsB. ThermodynamicsC. OpticsD. Quantum mechanics。
海洋放线菌
Contents 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actinomycetes in the marine environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Role of actinomycetes in marine environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rare actinomycetes and selective isolation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molecular approaches to search for indigenous marine actinomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Different genera of marine actinomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marine streptomycetes – a boundary microorganism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fermentation process for metabolites production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Secondary metabolites from actinomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Novel/new metabolites from marine actinomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 00 00 00 00 00 00 00 00 00 00 00 00 00
2005-A global Malmquist productivity index
A global Malmquist productivity indexJesu ´s T.Pastor a ,C.A.Knox Lovell b ,TaCentro de Investigacio ´n Operativa,Universidad Miguel Herna ´ndez,03206Elche (Alicante),SpainbDepartment of Economics,University of Georgia,Athens,GA 30602,USA Received 2June 2004;received in revised form 24January 2005;accepted 16February 2005Available online 23May 2005AbstractThe geometric mean Malmquist productivity index is not circular,and its adjacent period components can provide different measures of productivity change.We propose a global Malmquist productivity index that is circular,and that gives a single measure of productivity change.D 2005Elsevier B.V .All rights reserved.Keywords:Malmquist productivity index;Circularity JEL classification:C43;D24;O471.IntroductionThe geometric mean form of the contemporaneous Malmquist productivity index,introduced by Caves et al.(1982),is not circular.Whether this is a serious problem depends on the powers of persuasion of Fisher (1922),who dismissed the test,and Frisch (1936),who endorsed it.The index averages two possibly disparate measures of productivity change.Fa ¨re and Grosskopf (1996)state sufficient conditions on the adjacent period technologies for the index to satisfy circularity,and to average the same measures of productivity change.When linear programming techniques are used to compute and decompose the index,infeasibility can occur.Whether this is a serious problem depends on0165-1765/$-see front matter D 2005Elsevier B.V .All rights reserved.doi:10.1016/j.econlet.2005.02.013T Corresponding author.Tel.:+17065423689;fax:+17065423376.E-mail address:knox@ (C.A.K.Lovell).Economics Letters 88(2005)266–271/locate/econbasethe structure of the data.Xue and Harker(2002)provide necessary and sufficient conditions on the datafor LP infeasibility not to occur.We demonstrate that the source of all three problems is the specification of adjacent periodtechnologies in the construction of the index.We show that it is possible to specify a base periodtechnology in a way that solves all three problems,without having to impose restrictive conditions oneither the technologies or the data.Berg et al.(1992)proposed an index that compares adjacent period data using technology from a baseperiod.This index satisfies circularity and generates a single measure of productivity change,but it paysfor circularity with base period dependence,and it remains susceptible to LP infeasibility.Shestalova(2003)proposed an index having as its base a sequential technology formed from data ofall producers in all periods up to and including the two periods being compared.This index is immune toLP infeasibility,and it generates a single measure of productivity change,but it fails circularity and itprecludes technical regress.Thus no currently available Malmquist productivity index solves all three problems.We propose anew global index with technology formed from data of all producers in all periods.This index satisfiescircularity,it generates a single measure of productivity change,it allows technical regress,and it isimmune to LP infeasibility.In Section2we introduce and decompose the circular global index.Its efficiency change componentis the same as that of the contemporaneous index,but its technical change component is new.In Section3we relate it to the contemporaneous index.In Section4we provide an empirical illustration.Section5concludes.2.The global Malmquist productivity indexConsider a panel of i=1,...,I producers and t=1,...,T time periods.Producers use inputs x a R N+toproduce outputs y a R P+.We define two technologies.A contemporaneous benchmark technology isdefined as T c t={(x t,y t)|x t can produce y t}with k T c t=T c t,t=1,...,T,k N0.A global benchmarktechnology is defined as T c G=conv{T c1v...v T c T}.The subscript b c Q indicates that both benchmark technologies satisfy constant returns to scale.A contemporaneous Malmquist productivity index is defined on T c s asM scx t;y t;x tþ1;y tþ1ÀÁ¼D scx tþ1;y tþ1ðÞD scx t;y tðÞ;ð1Þwhere the output distance functions D c s(x,y)=min{/N0|(x,y//)a T c s},s=t,t+1.Since M c t(x t,y t,x t+1, y t+1)p M c t+1(x t,y t,x t+1,y t+1)without restrictions on the two technologies,the contemporaneous index is typically defined in geometric mean form as M c(x t,y t,x t+1,y t+1)=[M c t(x t,y t,x t+1,y t+1)ÂM c t+1(x t,y t,x t+1, y t+1)]1/2.A global Malmquist productivity index is defined on T c G asM Gcx t;y t;x tþ1;y tþ1ÀÁ¼D Gcx tþ1;y tþ1ðÞD Gcx t;y tðÞ;ð2Þwhere the output distance functions D c G(x,y)=min{/N0|(x,y//)a T c G}.J.T.Pastor,C.A.K.Lovell/Economics Letters88(2005)266–271267Both indexes compare (x t +1,y t +1)to (x t ,y t ),but they use different benchmarks.Since there is only one global benchmark technology,there is no need to resort to the geometric mean convention when defining the global index.M cGdecomposes as M G c x t ;y t ;x t þ1;f y t þ1ÀÁ¼D t þ1c x t þ1;y t þ1ðÞD t c x t ;y t ðÞÂD G c x t þ1;y t þ1ðÞD t þ1c x t þ1;y t þ1ðÞÂD t cx t ;y t ðÞD Gc x t ;y t ðÞ&'¼TE t þ1c x t þ1;y t þ1ðÞTE t c x t ;y t ðÞÂD G c Àx t þ1;y t þ1=D t þ1c x t þ1;y t þ1ðÞÁD G c x t ;y t =D t cx t ;y t ðÞÀÁ()¼EC c ÂBPG G ;t þ1cx t þ1;y t þ1ðÞBPG cx t ;y tðÞ()¼EC c ÂBPC c ;ð3Þwhere EC c is the usual efficiency change indicator and BPG c G,s V 1is a best practice gap between T c Gand T c s measured along rays (x s ,y s),s =t ,t +1.BPC c is the change in BPG c ,and provides a new measure of technical change.BPC c f 1indicates whether the benchmark technology in period t +1in the region[(x t +1,y t +1/D ct +1(x t +1,y t +1))]is closer to or farther away from the global benchmark technology than is the benchmark technology in period t in the region [(x t ,y t /D ct (x t ,y t ))].M c G has four virtues.First,like any fixed base index,M cGis circular,and since EC c is circular,so is BPC c .Second,each provides a single measure,with no need to take the geometric mean of disparate adjacent period measures.Third,but not shown here,the decomposition in (3)can be extended to generate a three-way decomposition that is structurally identical to the Ray and Desli (1997)decomposition of the contemporaneous index.M cGand M c share a common efficiency change component,but they have different technical change and scale components,and so M c Gp M c without restrictions on the technologies.Finally,the technical change and scale components of M c Gare immune to the LP infeasibility problem that plagues these components of M c .paring the global and contemporaneous indexes The ratioM G c =M c¼M G c =M t þ1cÀÁÂM G c =M t cÀÁÂÃ1=2¼D G cx t þ1;y t þ1=D t þ1c x t þ1;y t þ1ðÞÀÁD G c x t ;y t =D t þ1c x t ;y t ðÞÀÁ"#ÂD G c x t þ1;y t þ1=D t c x t þ1;y t þ1ðÞÀÁD G c x t ;y t =D t c x t ;y t ðÞÀÁ"#()1=2¼BPG G ;t þ1cx t þ1;y t þ1ðÞBPG G ;t þ1cx t ;y tðÞ"#ÂBPG G ;t c xt þ1;y t þ1ðÞBPG G ;t c x t ;y tðÞ"#()1=2ð4Þis the geometric mean of two terms,each being a ratio of benchmark technology gaps along differentrays.M c G /M c f 1as projections onto T c t and T c t +1of period t +1data are closer to,equidistant from,orfarther away from T c G than projections onto T c t and T ct +1of period t data are.J.T.Pastor,C.A.K.Lovell /Economics Letters 88(2005)266–271268J.T.Pastor,C.A.K.Lovell/Economics Letters88(2005)266–271269 Table1Electricity generation data,annual means1977198219871992 Output(000MW h)13,70013,86016,18017,270 Labor(#FTE)1373179719952021 Fuel(billion BTU)1288144116671824 Capital(To¨rnqvist)44,756211,622371,041396,386 M c G=M c if BPG c G,s(x t+1,y t+1)=BPG c G,s(x t,y t),s=t,t+1.From the first equality in(4),this condition is equivalent to the condition M c G=M c s,s=t,t+1.If this condition holds for all s,it is equivalent to the condition M c t=M c1for all t.Althin(2001)has shown that a sufficient condition for base period independence is that technical change be Hicks output-neutral(HON).Hence HON is also sufficient for M c G=M c.4.An empirical illustrationWe summarize an application intended to illustrate the behavior of M c G,and to compare its performance with that of M c.We analyze a panel of93US electricity generating firms in four years (1977,1982,1997,1992).The firms use labor(FTE employees),fuel(BTUs of energy)and capital(a multilateral To¨rnqvist index)to generate electricity(net generation in MW h).The data are summarized in Table1.Electricity generation increased by proportionately less than each input did.The main cause of the rapid increase in the capital input was the enactment of environmental regulations mandating the installation of pollution abatement equipment.We are unable to disaggregate the capital input into its productive and abatement components.Empirical findings are summarized in Table2.The first three rows report decomposition(3)of M c G, and the final three rows report M c and its two adjacent period components.Columns correspond to time periods.M c G shows a large productivity decline from1977to1982,followed by weak productivity growth. Cumulative productivity in1992was25%lower than in1977.M c G calculated using1992and1977data generates the same value,verifying that it is circular.The efficiency change component EC c of M c G(and M c)is also circular,and cumulates to an18% improvement.Best practice change,BPC c,is also circular,and declined by35%.Capital investment in Table2Global and contemporaneous Malmquist productivity indexes1977–19821982–19871987–1992Cumulative productivity1977–1992 M c G0.685 1.064 1.0390.7570.757EC c 1.163 1.0890.929 1.176 1.176 BPC c0.5890.977 1.1180.6440.644M c0.4310.895 1.0390.4000.592M c t0.7130.902 1.0530.678 1.333M c t+10.2600.887 1.0240.2360.263pollution abatement equipment generated cleaner air but not more electricity.Consequently catching up with deteriorating best practice was relatively easy.Turning to the contemporaneous index M c reported in the final three rows,the story is not so clear.Cumulative productivity in 1992was 60%lower than in 1977.However calculating M c using 1992and 1977data generates a smaller 40%decline,verifying that M c is not circular.Neither figure is close to the25%decline reported by M cG,verifying that technical change was not HON,but (pollution abatement)capital-using.The lack of circularity is reflected in the frequently large differences between M ct and M c t +1,which give conflicting signals when computed using 1992and 1977data,with M c tsignaling productivitygrowth and M ct +1signaling productivity decline.Although not reported in Table 2,we have calculated three-way decompositions of M cG and M c .All three components of M c G are circular,and LP infeasibility does not occur.In contrast,the technical change and scale components of M c are not circular,and infeasibility occurs for 13observations.The circular global index M cGtells a single story about productivity change,and its decomposition is intuitively appealing in light of what we know about the industry during the cking circularity,M c and its two adjacent period components tell different stories that are often contradictory.Thedifferences between M cGand M c are a consequence of the capital-using bias of technical change,which was regressive due to the mandated installation of pollution abatement equipment,augmented perhaps by the rate base padding that was prevalent during the period.5.ConclusionsThe contemporaneous Malmquist productivity index is not circular,its adjacent period components can give conflicting signals,and it is susceptible to LP infeasibility.The global Malmquist productivity index and each of its components is circular,it provides single measures of productivity change and its components,and it is immune to LP infeasibility.The global index decomposes into the same sources of productivity change as the contemporaneous index does.A sufficient condition for equality of the two indexes,and their respective components,is Hicks output neutrality of technical change.The global index must be recomputed when a new time period is incorporated.Diewert’s (1987)assertion that b ...economic history has to be rewritten ...Q when new data are incorporated is the base period dependency problem revisited.The problem can be serious when using base periods t =1and t =T ,but it is likely to be benign when using global base periods {1,...,T }and {1,...,T +1}.While new data may change the global frontier,the rewriting of history is likely to be quantitative rather than qualitative.ReferencesAlthin,R.,2001.Measurement of productivity changes:two Malmquist index approaches.Journal of Productivity Analysis 16,107–128.Berg,S.A.,Førsund,F.R.,Jansen,E.S.,1992.Malmquist indices of productivity growth during the deregulation of Norwegian banking,1980–89.Scandinavian Journal of Economics 94,211–228(Supplement).Caves,D.W.,Christensen,L.R.,Diewert,W.E.,1982.The economic theory of index numbers and the measurement of input output,and productivity.Econometrica 50,1393–1414.J.T.Pastor,C.A.K.Lovell /Economics Letters 88(2005)266–271270J.T.Pastor,C.A.K.Lovell/Economics Letters88(2005)266–271271 Diewert,W.E.,1987.Index numbers.In:Eatwell,J.,Milgate,M.,Newman,P.(Eds.),The New Palgrave:A Dictionary of Economics,vol.2.The Macmillan Press,New York.Fa¨re,R.,Grosskopf,S.,1996.Intertemporal Production Frontiers:With Dynamic DEA.Kluwer Academic Publishers,Boston. Fisher,I.,1922.The Making of Index Numbers.Houghton Mifflin,Boston.Frisch,R.,1936.Annual survey of general economic theory:the problem of index numbers.Econometrica4,1–38.Ray,S.C.,Desli,E.,1997.Productivity growth,technical progress,and efficiency change in industrialized countries:comment.American Economic Review87,1033–1039.Shestalova,V.,2003.Sequential Malmquist indices of productivity growth:an application to OECD industrial activities.Journal of Productivity Analysis19,211–226.Xue,M.,Harker,P.T.,2002.Note:ranking DMUs with infeasible super-efficiency in DEA models.Management Science48, 705–710.。
专八英语阅读
英语专业八级考试TEM-8阅读理解练习册(1)(英语专业2012级)UNIT 1Text AEvery minute of every day, what ecologist生态学家James Carlton calls a global ―conveyor belt‖, redistributes ocean organisms生物.It’s planetwide biological disruption生物的破坏that scientists have barely begun to understand.Dr. Carlton —an oceanographer at Williams College in Williamstown,Mass.—explains that, at any given moment, ―There are several thousand marine species traveling… in the ballast water of ships.‖ These creatures move from coastal waters where they fit into the local web of life to places where some of them could tear that web apart. This is the larger dimension of the infamous无耻的,邪恶的invasion of fish-destroying, pipe-clogging zebra mussels有斑马纹的贻贝.Such voracious贪婪的invaders at least make their presence known. What concerns Carlton and his fellow marine ecologists is the lack of knowledge about the hundreds of alien invaders that quietly enter coastal waters around the world every day. Many of them probably just die out. Some benignly亲切地,仁慈地—or even beneficially — join the local scene. But some will make trouble.In one sense, this is an old story. Organisms have ridden ships for centuries. They have clung to hulls and come along with cargo. What’s new is the scale and speed of the migrations made possible by the massive volume of ship-ballast water压载水— taken in to provide ship stability—continuously moving around the world…Ships load up with ballast water and its inhabitants in coastal waters of one port and dump the ballast in another port that may be thousands of kilometers away. A single load can run to hundreds of gallons. Some larger ships take on as much as 40 million gallons. The creatures that come along tend to be in their larva free-floating stage. When discharged排出in alien waters they can mature into crabs, jellyfish水母, slugs鼻涕虫,蛞蝓, and many other forms.Since the problem involves coastal species, simply banning ballast dumps in coastal waters would, in theory, solve it. Coastal organisms in ballast water that is flushed into midocean would not survive. Such a ban has worked for North American Inland Waterway. But it would be hard to enforce it worldwide. Heating ballast water or straining it should also halt the species spread. But before any such worldwide regulations were imposed, scientists would need a clearer view of what is going on.The continuous shuffling洗牌of marine organisms has changed the biology of the sea on a global scale. It can have devastating effects as in the case of the American comb jellyfish that recently invaded the Black Sea. It has destroyed that sea’s anchovy鳀鱼fishery by eating anchovy eggs. It may soon spread to western and northern European waters.The maritime nations that created the biological ―conveyor belt‖ should support a coordinated international effort to find out what is going on and what should be done about it. (456 words)1.According to Dr. Carlton, ocean organism‟s are_______.A.being moved to new environmentsB.destroying the planetC.succumbing to the zebra musselD.developing alien characteristics2.Oceanographers海洋学家are concerned because_________.A.their knowledge of this phenomenon is limitedB.they believe the oceans are dyingC.they fear an invasion from outer-spaceD.they have identified thousands of alien webs3.According to marine ecologists, transplanted marinespecies____________.A.may upset the ecosystems of coastal watersB.are all compatible with one anotherC.can only survive in their home watersD.sometimes disrupt shipping lanes4.The identified cause of the problem is_______.A.the rapidity with which larvae matureB. a common practice of the shipping industryC. a centuries old speciesD.the world wide movement of ocean currents5.The article suggests that a solution to the problem__________.A.is unlikely to be identifiedB.must precede further researchC.is hypothetically假设地,假想地easyD.will limit global shippingText BNew …Endangered‟ List Targets Many US RiversIt is hard to think of a major natural resource or pollution issue in North America today that does not affect rivers.Farm chemical runoff残渣, industrial waste, urban storm sewers, sewage treatment, mining, logging, grazing放牧,military bases, residential and business development, hydropower水力发电,loss of wetlands. The list goes on.Legislation like the Clean Water Act and Wild and Scenic Rivers Act have provided some protection, but threats continue.The Environmental Protection Agency (EPA) reported yesterday that an assessment of 642,000 miles of rivers and streams showed 34 percent in less than good condition. In a major study of the Clean Water Act, the Natural Resources Defense Council last fall reported that poison runoff impairs损害more than 125,000 miles of rivers.More recently, the NRDC and Izaak Walton League warned that pollution and loss of wetlands—made worse by last year’s flooding—is degrading恶化the Mississippi River ecosystem.On Tuesday, the conservation group保护组织American Rivers issued its annual list of 10 ―endangered‖ and 20 ―threatened‖ rivers in 32 states, the District of Colombia, and Canada.At the top of the list is the Clarks Fork of the Yellowstone River, whereCanadian mining firms plan to build a 74-acre英亩reservoir水库,蓄水池as part of a gold mine less than three miles from Yellowstone National Park. The reservoir would hold the runoff from the sulfuric acid 硫酸used to extract gold from crushed rock.―In the event this tailings pond failed, the impact to th e greater Yellowstone ecosystem would be cataclysmic大变动的,灾难性的and the damage irreversible不可逆转的.‖ Sen. Max Baucus of Montana, chairman of the Environment and Public Works Committee, wrote to Noranda Minerals Inc., an owner of the ― New World Mine‖.Last fall, an EPA official expressed concern about the mine and its potential impact, especially the plastic-lined storage reservoir. ― I am unaware of any studies evaluating how a tailings pond尾矿池,残渣池could be maintained to ensure its structural integrity forev er,‖ said Stephen Hoffman, chief of the EPA’s Mining Waste Section. ―It is my opinion that underwater disposal of tailings at New World may present a potentially significant threat to human health and the environment.‖The results of an environmental-impact statement, now being drafted by the Forest Service and Montana Department of State Lands, could determine the mine’s future…In its recent proposal to reauthorize the Clean Water Act, the Clinton administration noted ―dramatically improved water quality since 1972,‖ when the act was passed. But it also reported that 30 percent of riverscontinue to be degraded, mainly by silt泥沙and nutrients from farm and urban runoff, combined sewer overflows, and municipal sewage城市污水. Bottom sediments沉积物are contaminated污染in more than 1,000 waterways, the administration reported in releasing its proposal in January. Between 60 and 80 percent of riparian corridors (riverbank lands) have been degraded.As with endangered species and their habitats in forests and deserts, the complexity of ecosystems is seen in rivers and the effects of development----beyond the obvious threats of industrial pollution, municipal waste, and in-stream diversions改道to slake消除the thirst of new communities in dry regions like the Southwes t…While there are many political hurdles障碍ahead, reauthorization of the Clean Water Act this year holds promise for US rivers. Rep. Norm Mineta of California, who chairs the House Committee overseeing the bill, calls it ―probably the most important env ironmental legislation this Congress will enact.‖ (553 words)6.According to the passage, the Clean Water Act______.A.has been ineffectiveB.will definitely be renewedC.has never been evaluatedD.was enacted some 30 years ago7.“Endangered” rivers are _________.A.catalogued annuallyB.less polluted than ―threatened rivers‖C.caused by floodingD.adjacent to large cities8.The “cataclysmic” event referred to in paragraph eight would be__________.A. fortuitous偶然的,意外的B. adventitious外加的,偶然的C. catastrophicD. precarious不稳定的,危险的9. The owners of the New World Mine appear to be______.A. ecologically aware of the impact of miningB. determined to construct a safe tailings pondC. indifferent to the concerns voiced by the EPAD. willing to relocate operations10. The passage conveys the impression that_______.A. Canadians are disinterested in natural resourcesB. private and public environmental groups aboundC. river banks are erodingD. the majority of US rivers are in poor conditionText CA classic series of experiments to determine the effects ofoverpopulation on communities of rats was reported in February of 1962 in an article in Scientific American. The experiments were conducted by a psychologist, John B. Calhoun and his associates. In each of these experiments, an equal number of male and female adult rats were placed in an enclosure and given an adequate supply of food, water, and other necessities. The rat populations were allowed to increase. Calhoun knew from experience approximately how many rats could live in the enclosures without experiencing stress due to overcrowding. He allowed the population to increase to approximately twice this number. Then he stabilized the population by removing offspring that were not dependent on their mothers. He and his associates then carefully observed and recorded behavior in these overpopulated communities. At the end of their experiments, Calhoun and his associates were able to conclude that overcrowding causes a breakdown in the normal social relationships among rats, a kind of social disease. The rats in the experiments did not follow the same patterns of behavior as rats would in a community without overcrowding.The females in the rat population were the most seriously affected by the high population density: They showed deviant异常的maternal behavior; they did not behave as mother rats normally do. In fact, many of the pups幼兽,幼崽, as rat babies are called, died as a result of poor maternal care. For example, mothers sometimes abandoned their pups,and, without their mothers' care, the pups died. Under normal conditions, a mother rat would not leave her pups alone to die. However, the experiments verified that in overpopulated communities, mother rats do not behave normally. Their behavior may be considered pathologically 病理上,病理学地diseased.The dominant males in the rat population were the least affected by overpopulation. Each of these strong males claimed an area of the enclosure as his own. Therefore, these individuals did not experience the overcrowding in the same way as the other rats did. The fact that the dominant males had adequate space in which to live may explain why they were not as seriously affected by overpopulation as the other rats. However, dominant males did behave pathologically at times. Their antisocial behavior consisted of attacks on weaker male,female, and immature rats. This deviant behavior showed that even though the dominant males had enough living space, they too were affected by the general overcrowding in the enclosure.Non-dominant males in the experimental rat communities also exhibited deviant social behavior. Some withdrew completely; they moved very little and ate and drank at times when the other rats were sleeping in order to avoid contact with them. Other non-dominant males were hyperactive; they were much more active than is normal, chasing other rats and fighting each other. This segment of the rat population, likeall the other parts, was affected by the overpopulation.The behavior of the non-dominant males and of the other components of the rat population has parallels in human behavior. People in densely populated areas exhibit deviant behavior similar to that of the rats in Calhoun's experiments. In large urban areas such as New York City, London, Mexican City, and Cairo, there are abandoned children. There are cruel, powerful individuals, both men and women. There are also people who withdraw and people who become hyperactive. The quantity of other forms of social pathology such as murder, rape, and robbery also frequently occur in densely populated human communities. Is the principal cause of these disorders overpopulation? Calhoun’s experiments suggest that it might be. In any case, social scientists and city planners have been influenced by the results of this series of experiments.11. Paragraph l is organized according to__________.A. reasonsB. descriptionC. examplesD. definition12.Calhoun stabilized the rat population_________.A. when it was double the number that could live in the enclosure without stressB. by removing young ratsC. at a constant number of adult rats in the enclosureD. all of the above are correct13.W hich of the following inferences CANNOT be made from theinformation inPara. 1?A. Calhoun's experiment is still considered important today.B. Overpopulation causes pathological behavior in rat populations.C. Stress does not occur in rat communities unless there is overcrowding.D. Calhoun had experimented with rats before.14. Which of the following behavior didn‟t happen in this experiment?A. All the male rats exhibited pathological behavior.B. Mother rats abandoned their pups.C. Female rats showed deviant maternal behavior.D. Mother rats left their rat babies alone.15. The main idea of the paragraph three is that __________.A. dominant males had adequate living spaceB. dominant males were not as seriously affected by overcrowding as the otherratsC. dominant males attacked weaker ratsD. the strongest males are always able to adapt to bad conditionsText DThe first mention of slavery in the statutes法令,法规of the English colonies of North America does not occur until after 1660—some forty years after the importation of the first Black people. Lest we think that existed in fact before it did in law, Oscar and Mary Handlin assure us, that the status of B lack people down to the 1660’s was that of servants. A critique批判of the Handlins’ interpretation of why legal slavery did not appear until the 1660’s suggests that assumptions about the relation between slavery and racial prejudice should be reexamined, and that explanation for the different treatment of Black slaves in North and South America should be expanded.The Handlins explain the appearance of legal slavery by arguing that, during the 1660’s, the position of white servants was improving relative to that of black servants. Thus, the Handlins contend, Black and White servants, heretofore treated alike, each attained a different status. There are, however, important objections to this argument. First, the Handlins cannot adequately demonstrate that t he White servant’s position was improving, during and after the 1660’s; several acts of the Maryland and Virginia legislatures indicate otherwise. Another flaw in the Handlins’ interpretation is their assumption that prior to the establishment of legal slavery there was no discrimination against Black people. It is true that before the 1660’s Black people were rarely called slaves. But this shouldnot overshadow evidence from the 1630’s on that points to racial discrimination without using the term slavery. Such discrimination sometimes stopped short of lifetime servitude or inherited status—the two attributes of true slavery—yet in other cases it included both. The Handlins’ argument excludes the real possibility that Black people in the English colonies were never treated as the equals of White people.The possibility has important ramifications后果,影响.If from the outset Black people were discriminated against, then legal slavery should be viewed as a reflection and an extension of racial prejudice rather than, as many historians including the Handlins have argued, the cause of prejudice. In addition, the existence of discrimination before the advent of legal slavery offers a further explanation for the harsher treatment of Black slaves in North than in South America. Freyre and Tannenbaum have rightly argued that the lack of certain traditions in North America—such as a Roman conception of slavery and a Roman Catholic emphasis on equality— explains why the treatment of Black slaves was more severe there than in the Spanish and Portuguese colonies of South America. But this cannot be the whole explanation since it is merely negative, based only on a lack of something. A more compelling令人信服的explanation is that the early and sometimes extreme racial discrimination in the English colonies helped determine the particular nature of the slavery that followed. (462 words)16. Which of the following is the most logical inference to be drawn from the passage about the effects of “several acts of the Maryland and Virginia legislatures” (Para.2) passed during and after the 1660‟s?A. The acts negatively affected the pre-1660’s position of Black as wellas of White servants.B. The acts had the effect of impairing rather than improving theposition of White servants relative to what it had been before the 1660’s.C. The acts had a different effect on the position of white servants thandid many of the acts passed during this time by the legislatures of other colonies.D. The acts, at the very least, caused the position of White servants toremain no better than it had been before the 1660’s.17. With which of the following statements regarding the status ofBlack people in the English colonies of North America before the 1660‟s would the author be LEAST likely to agree?A. Although black people were not legally considered to be slaves,they were often called slaves.B. Although subject to some discrimination, black people had a higherlegal status than they did after the 1660’s.C. Although sometimes subject to lifetime servitude, black peoplewere not legally considered to be slaves.D. Although often not treated the same as White people, black people,like many white people, possessed the legal status of servants.18. According to the passage, the Handlins have argued which of thefollowing about the relationship between racial prejudice and the institution of legal slavery in the English colonies of North America?A. Racial prejudice and the institution of slavery arose simultaneously.B. Racial prejudice most often the form of the imposition of inheritedstatus, one of the attributes of slavery.C. The source of racial prejudice was the institution of slavery.D. Because of the influence of the Roman Catholic Church, racialprejudice sometimes did not result in slavery.19. The passage suggests that the existence of a Roman conception ofslavery in Spanish and Portuguese colonies had the effect of _________.A. extending rather than causing racial prejudice in these coloniesB. hastening the legalization of slavery in these colonies.C. mitigating some of the conditions of slavery for black people in these coloniesD. delaying the introduction of slavery into the English colonies20. The author considers the explanation put forward by Freyre andTannenbaum for the treatment accorded B lack slaves in the English colonies of North America to be _____________.A. ambitious but misguidedB. valid有根据的but limitedC. popular but suspectD. anachronistic过时的,时代错误的and controversialUNIT 2Text AThe sea lay like an unbroken mirror all around the pine-girt, lonely shores of Orr’s Island. Tall, kingly spruce s wore their regal王室的crowns of cones high in air, sparkling with diamonds of clear exuded gum流出的树胶; vast old hemlocks铁杉of primeval原始的growth stood darkling in their forest shadows, their branches hung with long hoary moss久远的青苔;while feathery larches羽毛般的落叶松,turned to brilliant gold by autumn frosts, lighted up the darker shadows of the evergreens. It was one of those hazy朦胧的, calm, dissolving days of Indian summer, when everything is so quiet that the fainest kiss of the wave on the beach can be heard, and white clouds seem to faint into the blue of the sky, and soft swathing一长条bands of violet vapor make all earth look dreamy, and give to the sharp, clear-cut outlines of the northern landscape all those mysteries of light and shade which impart such tenderness to Italian scenery.The funeral was over,--- the tread鞋底的花纹/ 踏of many feet, bearing the heavy burden of two broken lives, had been to the lonely graveyard, and had come back again,--- each footstep lighter and more unconstrained不受拘束的as each one went his way from the great old tragedy of Death to the common cheerful of Life.The solemn black clock stood swaying with its eternal ―tick-tock, tick-tock,‖ in the kitchen of the brown house on Orr’s Island. There was there that sense of a stillness that can be felt,---such as settles down on a dwelling住处when any of its inmates have passed through its doors for the last time, to go whence they shall not return. The best room was shut up and darkened, with only so much light as could fall through a little heart-shaped hole in the window-shutter,---for except on solemn visits, or prayer-meetings or weddings, or funerals, that room formed no part of the daily family scenery.The kitchen was clean and ample, hearth灶台, and oven on one side, and rows of old-fashioned splint-bottomed chairs against the wall. A table scoured to snowy whiteness, and a little work-stand whereon lay the Bible, the Missionary Herald, and the Weekly Christian Mirror, before named, formed the principal furniture. One feature, however, must not be forgotten, ---a great sea-chest水手用的储物箱,which had been the companion of Zephaniah through all the countries of the earth. Old, and battered破旧的,磨损的, and unsightly难看的it looked, yet report said that there was good store within which men for the most part respect more than anything else; and, indeed it proved often when a deed of grace was to be done--- when a woman was suddenly made a widow in a coast gale大风,狂风, or a fishing-smack小渔船was run down in the fogs off the banks, leaving in some neighboring cottage a family of orphans,---in all such cases, the opening of this sea-chest was an event of good omen 预兆to the bereaved丧亲者;for Zephaniah had a large heart and a large hand, and was apt有…的倾向to take it out full of silver dollars when once it went in. So the ark of the covenant约柜could not have been looked on with more reverence崇敬than the neighbours usually showed to Captain Pennel’s sea-chest.1. The author describes Orr‟s Island in a(n)______way.A.emotionally appealing, imaginativeB.rational, logically preciseC.factually detailed, objectiveD.vague, uncertain2.According to the passage, the “best room”_____.A.has its many windows boarded upB.has had the furniture removedC.is used only on formal and ceremonious occasionsD.is the busiest room in the house3.From the description of the kitchen we can infer that thehouse belongs to people who_____.A.never have guestsB.like modern appliancesC.are probably religiousD.dislike housework4.The passage implies that_______.A.few people attended the funeralB.fishing is a secure vocationC.the island is densely populatedD.the house belonged to the deceased5.From the description of Zephaniah we can see thathe_________.A.was physically a very big manB.preferred the lonely life of a sailorC.always stayed at homeD.was frugal and saved a lotText BBasic to any understanding of Canada in the 20 years after the Second World War is the country' s impressive population growth. For every three Canadians in 1945, there were over five in 1966. In September 1966 Canada's population passed the 20 million mark. Most of this surging growth came from natural increase. The depression of the 1930s and the war had held back marriages, and the catching-up process began after 1945. The baby boom continued through the decade of the 1950s, producing a population increase of nearly fifteen percent in the five years from 1951 to 1956. This rate of increase had been exceeded only once before in Canada's history, in the decade before 1911 when the prairies were being settled. Undoubtedly, the good economic conditions of the 1950s supported a growth in the population, but the expansion also derived from a trend toward earlier marriages and an increase in the average size of families; In 1957 the Canadian birth rate stood at 28 per thousand, one of the highest in the world. After the peak year of 1957, thebirth rate in Canada began to decline. It continued falling until in 1966 it stood at the lowest level in 25 years. Partly this decline reflected the low level of births during the depression and the war, but it was also caused by changes in Canadian society. Young people were staying at school longer, more women were working; young married couples were buying automobiles or houses before starting families; rising living standards were cutting down the size of families. It appeared that Canada was once more falling in step with the trend toward smaller families that had occurred all through theWestern world since the time of the Industrial Revolution. Although the growth in Canada’s population had slowed down by 1966 (the cent), another increase in the first half of the 1960s was only nine percent), another large population wave was coming over the horizon. It would be composed of the children of the children who were born during the period of the high birth rate prior to 1957.6. What does the passage mainly discuss?A. Educational changes in Canadian society.B. Canada during the Second World War.C. Population trends in postwar Canada.D. Standards of living in Canada.7. According to the passage, when did Canada's baby boom begin?A. In the decade after 1911.B. After 1945.C. During the depression of the 1930s.D. In 1966.8. The author suggests that in Canada during the 1950s____________.A. the urban population decreased rapidlyB. fewer people marriedC. economic conditions were poorD. the birth rate was very high9. When was the birth rate in Canada at its lowest postwar level?A. 1966.B. 1957.C. 1956.D. 1951.10. The author mentions all of the following as causes of declines inpopulation growth after 1957 EXCEPT_________________.A. people being better educatedB. people getting married earlierC. better standards of livingD. couples buying houses11.I t can be inferred from the passage that before the IndustrialRevolution_______________.A. families were largerB. population statistics were unreliableC. the population grew steadilyD. economic conditions were badText CI was just a boy when my father brought me to Harlem for the first time, almost 50 years ago. We stayed at the hotel Theresa, a grand brick structure at 125th Street and Seventh avenue. Once, in the hotel restaurant, my father pointed out Joe Louis. He even got Mr. Brown, the hotel manager, to introduce me to him, a bit punchy强力的but still champ焦急as fast as I was concerned.Much has changed since then. Business and real estate are booming. Some say a new renaissance is under way. Others decry责难what they see as outside forces running roughshod肆意践踏over the old Harlem. New York meant Harlem to me, and as a young man I visited it whenever I could. But many of my old haunts are gone. The Theresa shut down in 1966. National chains that once ignored Harlem now anticipate yuppie money and want pieces of this prime Manhattan real estate. So here I am on a hot August afternoon, sitting in a Starbucks that two years ago opened a block away from the Theresa, snatching抓取,攫取at memories between sips of high-priced coffee. I am about to open up a piece of the old Harlem---the New York Amsterdam News---when a tourist。
遗传学中英文词汇
中英文词汇--------------------------------------------------------------------------------近端着丝粒染色体(Acrocentric chromosome)——着丝粒靠近染色体端部的染色体。
加和原则(Additivity principle)——如果两个事件相互排斥,那么获得其中一个或另一个的概率为它们的各自概率之和。
等位基因(Allele)——在一既定基因座上一个基因的替换形式。
等位基因特异性寡核苷酸(Allele-specific oligonucleotide,ASO)——设计合成的寡核苷酸,可在适当条件下与特异序列杂交而不与其相关的序列杂交。
用针对每个等位基因序列设计的ASO甚至可容易地检出单个核苷酸的变异。
在几种设计相似、用来区分密切相关等位基因的方法中,ASO还可用作PCR引物。
等位基因异质性(Allelic heterogeneity)——在同一遗传基因座上,由不同的突变等位基因引起的相同或相似的表型。
α1-抗胰蛋白酶(α1-Antitrypsin)——是抑制弹性蛋白酶活性的一种丝氨酸蛋白酶抑制剂,该抑制剂的缺乏(如α1-抗胰蛋白酶不足)将导致严重的慢性肺和肝脏疾病。
Alu重复序列(Alu repetitive sequence)——位于基因间或内含子DNA中的中等重复序列,含有限制性内切酶AluⅠ的识别位点,这些序列长约300bp,并在人类基因组中重复出现约500,000次。
羊膜穿刺术(Amniocentesis)——一种产前诊断的方法,通常在妊娠4至6月抽取羊膜囊内婴儿四周的羊水进行。
扩增(Amplification)——一段DNA序列多个拷贝的产生。
非整倍体(Aneuploid)——指单倍体非整倍数的任何染色体数目。
通常非整倍体是指单条染色体的额外拷贝(三体性),或缺少单条染色体(单体性)。
由减数分裂或有丝分裂过程中染色体不分离所致。
抗病毒种质创制 项目指南
抗病毒种质创制项目指南英文回答:Antiviral germplasm creation is a vital aspect of virus resistance in plants. Developing new plant varieties with enhanced resistance to viral infections is crucial for ensuring food security and sustainable agriculture. In this project guide, I will outline the key steps and considerations involved in creating antiviral germplasm.1. Understanding the Virus: The first step in creating antiviral germplasm is to gain a comprehensiveunderstanding of the virus or viruses that the plant is susceptible to. This involves studying the virus's structure, replication cycle, modes of transmission, and interaction with the host plant. By understanding the virus, we can identify potential targets for resistance.2. Identifying Resistance Genes: Once we have a good understanding of the virus, the next step is to identifyand isolate resistance genes from other plant species that are known to be resistant to the virus. These resistance genes can be transferred to the susceptible plant through traditional breeding methods or genetic engineering techniques.3. Incorporating Resistance Genes: The identified resistance genes need to be incorporated into the susceptible plant's genome. This can be achieved through crossbreeding with resistant varieties or by using genetic engineering techniques such as gene editing or transformation. The goal is to introduce the resistance genes into the susceptible plant's genome while maintaining its desirable traits.4. Screening and Selection: After incorporating the resistance genes, the resulting plant populations need to be screened and selected for resistance. This involves exposing the plants to the virus and observing their response. Plants that show a high level of resistance are selected for further breeding or propagation.5. Field Testing and Evaluation: Selected resistant plants are then subjected to field testing to evaluatetheir performance under natural conditions. This helps determine their suitability for commercial cultivation. Factors such as yield, quality, and agronomic traits are assessed to ensure that the new varieties meet the requirements of farmers and consumers.6. Multiplication and Distribution: Once the resistant plant varieties have been successfully developed and evaluated, they need to be multiplied and distributed to farmers and growers. This involves producing sufficient quantities of seeds or plant materials for widespread adoption.中文回答:抗病毒种质创制是植物抗病毒能力的重要方面。
植物生理学——精选推荐
1:植物生理学有哪些主要的研究领域?细胞生理,代谢生理,生长发育生理,逆境生理。
2; 植物生理学与农林业生产实践有何相互关系?生产实践的需要是植物生理学产生的先决条件,也是植物生理学应用的必然归宿。
农林业生产进步的根本保证是依靠科学技术。
合理利用植物生理学的知识一实现农林业生产的高效、优质、高产的目标更为重要。
植物生理学与农林业生产的实践之间是一种紧密的互动关系。
植物生理学的发展能够推动农林业生产的进步,植物生理学在解决实际问题的时又会发现新的课题而推动植物生理学研究。
事实上,植物生理学在长期服务于农林业生产的实践中已经有了大量成功的先例。
3.植物生理学对农业生产的指导作用可表现在哪些方面?答:“植物生理学是合理农业的基础”,植物矿质营养学说的创立为无机肥料的施用奠定了理论基础,由于化肥的大量施用促使世界粮食产量急增;植物激素的陆续发现导致了植物生长调节剂和除草剂的普通应用,给农业生产带来革命性的变革;在光合作用与产量关系的理论指导下,植物生理学家与育种学家相结合,开创了以培育矮杆、耐肥、抗倒、叶片直立、株型紧凑的水稻、小麦品种为主要内容的“绿色革命”,使稻、麦产量获得了新的突破;植物细胞全能性理论的确立,不仅使人们掌握了如组织培养、细胞及原生质体培养等高效快速的植物无性繁殖新技术,而且为植物基因工程的开展和新种质的创造提供了先决条件。
按照我国农业生产发展的趋势,以下几方面可能成为今后植物生理学与农业生产相结合的重要研究领域:(1) 作物产量形成与高产理论。
(2) 环境生理与作物抗逆性。
(3) 设施农业中的作物生理学。
(4) 植物生理学与育种学相结合——作物生理育种等4、拟南芥有什么生物学和遗传学特征?我们为什么使用拟南芥做研究?常用拟南芥数据库有哪些?生物学特征:基生叶有柄呈莲座状,叶片倒卵形或匙形;茎生叶无柄,披针形或线形。
总状花序顶生,花瓣4片,白色,匙形。
长角果线形,长1~1.5厘米。
花期3~5月。
Nitrogen Cycling in the Ocean
10.1128/AEM.68.3.1015-1024.2002.2002, 68(3):1015. DOI:Appl. Environ. Microbiol. Jonathan P. Zehr and Bess B. WardPerspectives on Processes and ParadigmsNitrogen Cycling in the Ocean: New/content/68/3/1015Updated information and services can be found at: These include:REFERENCES/content/68/3/1015#ref-list-1at: This article cites 109 articles, 36 of which can be accessed free CONTENT ALERTSmore»articles cite this article), Receive: RSS Feeds, eTOCs, free email alerts (when new /site/misc/reprints.xhtml Information about commercial reprint orders: /site/subscriptions/To subscribe to to another ASM Journal go to: on November 14, 2014 by NANKAI UNIVERSITY/Downloaded fromA PPLIED AND E NVIRONMENTAL M ICROBIOLOGY,Mar.2002,p.1015–1024Vol.68,No.3 0099-2240/02/$04.00ϩ0DOI:10.1128/AEM.68.3.1015–1024.2002Copyright©2002,American Society for Microbiology.All Rights Reserved.MINIREVIEWNitrogen Cycling in the Ocean:New Perspectives on Processesand ParadigmsJonathan P.Zehr1*and Bess B.Ward2Department of Ocean Sciences,University of California,Santa Cruz,California95064,1and Department of Geosciences,Princeton University,Princeton,New Jersey085442The nitrogen(N)cycle is composed of multiple transforma-tions of nitrogenous compounds,catalyzed primarily by mi-crobes.The N cycle controls the availability of nitrogenous nutrients and biological productivity in marine systems(84) and thus is linked to thefixation of atmospheric carbon dioxide and export of carbon from the ocean’s surface(30).Human activities are influencing the N cycle even in the oceans(100), and some of the nitrogenous gaseous products of microbial metabolism are greenhouse gases that are potentially involved in controlling Earth’s climate.The last decade has brought the discovery of several new links and changes in our understanding of components of the marine N cycle.Much of our basic information about the N cycle is derived from measurements of transformation rates or from experiments with cultivated isolates.Generalization from the behavior and physiology of cultivated isolates can be mis-leading,since it appears that many marine microorganisms in situ have yet to be obtained in culture(33).The rapid increase in knowledge of genes and molecular biology has had an enor-mous impact on our understanding of the N cycle by making it possible to study the ecological underpinnings and diversity of microorganisms involved in specific N cycle components.For example,nitrate assimilation by photosynthetic picoplankton was assumed to be universal but has now been shown to be absent from some of the most abundant photosynthetic organ-isms on the planet.Conversely,nitrate assimilation by hetero-trophic bacteria was largely ignored,but it has recently been shown by both gene probes and physiological experiments that this capability is widespread in bacteria.Genetic and biochem-ical investigations have also changed our understanding of processes such as nitrification and denitrification,which were thought to be restricted to very specific habitats and microbes but in fact are more widely distributed.The N cycle is composed of oxidation-reduction reactions, many of which are used in the energy metabolism of microbes. Specific enzymes catalyze many of these reactions,and the enzymes and genes are useful targets for studying microbial processes such as assimilatory nitrate reduction,dissimilatory nitrate reduction,and N2fixation(Table1).Knowledge of the genes encoding enzymes involved in biogeochemical transfor-mations provides useful tools not only for assaying gene ex-pression but also for determining the diversity of microorgan-isms involved in specific N cycle transformations.We still know very little about the link between the ecology of the N cycle and the redundancy of microbes and genes in the marine environ-ment.Understanding these links is important for determining the role of microbial diversity in ecosystem processes and the sensitivity of the environment to perturbations.Regulatory proteins are also useful for investigating natural populations for“N status,”the degree of cellular N deficiency or suffi-ciency.In the following sections,we review some important aspects of the N cycle that have undergone major reevaluation in recent years.Rather than present a detailed review of the entire cycle,we focus on important recent changes in our understanding of these aspects and specific components of the cycle.Understanding these features of the nitrogen cycle is critical for understanding the global biogeochemical cycles. Here we focus on the microbiological underpinnings of these processes;the global implications for the biogeochemical cy-cling of nitrogen are beyond the scope of this review.INORGANIC AND ORGANIC N UPTAKEIt is usually assumed that most microorganisms can use inorganic N in the form of nitrate,nitrite,and ammonium. Particularly in the oligotrophic open ocean gyres,low concen-trations of these compounds(Ͻ0.03to0.1M[12])can limit the rate of productivity in the surface layer(0-to200-m depth), but N can regulate productivity even in coastal upwelling re-gions(55).In some regions,including coastal regions and the high-nutrient,low-chlorophyll regions,upwelling or runoff can supply N in concentrations that exceed phytoplankton demand. Thus,there are large geographical variations in the sources andfluxes of nitrate and ammonium.The geographic aspects of the marine N cycle are beyond the scope of this review but are covered elsewhere(12,16,103).Thefirst comprehensive view of the N cycle in the surface ocean proposed that inorganic N was taken up by phytoplank-ton and that the N was subsequently recycled from phytoplank-ton cells by heterotrophs,both large grazers(e.g.,planktonic invertebrates)and microbial decomposers(Fig.1,pathway A; Fig.2).The death,lysis,and decay of the phytoplankton,either after ingestion by“herbivores”or because of physiological stressors(such as nutrient limitation,temperature,or other*Corresponding author.Mailing address:Department of Ocean Sciences,Earth and Marine Sciences Building,Room A438,Universityof California,Santa Cruz,CA95064.Phone:(831)459-4009.Fax: (831)459-4882.E-mail:zehrj@.1015 on November 14, 2014 by NANKAI UNIVERSITY / Downloaded fromfactors),liberated N in the form of dissolved organic N or ammonium,which was termed“regenerated”N(Fig.1,path-way B;Fig.2)(28).Biological N2fixation,the reduction of atmospheric dinitrogen gas(N2)to ammonia(Fig.1,pathway C),was thought to be insignificant in the open ocean,and essentially all pelagic Nfixation was ascribed to two genera of N2-fixing microbes.Nitrate was believed to be supplied to the upper ocean primarily by mixing,advection and diffusion from deep ocean water,or terrestrial runoff(Fig.1,pathway D). Because nitrifying bacteria are inhibited by light,it was as-sumed that nitrification(Fig.1,pathway E)proceeded only in deep water(Fig.1,pathway E);therefore,the only source of nitrate in surface waters was water mixing from the deep ocean reservoir(Fig.1,pathway D).Nitrate,nitrite,and ammonium,called dissolved inorganic nitrogen(DIN),can be taken up(via membrane transporters) and assimilated by many microorganisms.Nitrate is assimilated after sequential reduction to nitrite(assimilatory nitrate reduc-tase)and ammonium(assimilatory nitrite reductase).Phyto-plankton(both eukaryotes and cyanobacteria),in general,pre-FIG.1.Conceptual diagram of major features of the nitrogen cycle in coastal shelf and upwelling(I),OMZs(II),surface waters of the openocean(III),and deep water(IV).PON,particulate organic nitrogen.Dashed lines indicate transformations involving multiple steps.Pathways:A,DIN assimilation;B,ammonium regeneration;C,nitrogenfixation;D,nitrate diffusion/advection from deep water;E,nitrification;F,nitrification;G,denitrification.TABLE1.Nitrogen cycle gene probesTransformation Gene(s)a Protein Reference(s)N2fixation nifHDK Nitrogenase117Nitrite assimilation nir Nitrite reductase102Nitrate assimilation narB,nasA Assimilatory nitrate reductase1Ammonium assimilation glnA Glutamine synthetase54Nitrate respiration and denitrification nirS Nitrate reductase8,18,87nirK Nitrite reductasenorB Nitric oxide reductasenosZ Nitrous oxide reductaseOrganic N metabolism ure Urease25Ammonium oxidation/nitrification amo Ammonia monooxygenase2,83Nitrogen regulation(cyanobacteria)ntcA Nitrogen regulatory protein58a Nitrogen cycle genes for which probes or PCR primers have been designed.1016MINIREVIEW A PPL.E NVIRON.M ICROBIOL.on November 14, 2014 by NANKAI UNIVERSITY /Downloaded fromfer ammonium to nitrate,presumably due to the additional energy and reductant necessary to reduce nitrate to ammo-nium.However,this preference is not universal,as the reduc-tant requirement for nitrate does not necessarily result in de-creased growth rate (95).Although it is still likely that most microorganisms prefer ammonium to nitrate,it has recently become clear that (i)not all phytoplankton use nitrate and (ii)some bacteria can use nitrate and some cannot.While it was previously assumed that eukaryotic phytoplankton dominated both photosynthesis and DIN assimilation,it is now known that two major groups of small unicellular cyanobacteria (the Syn-echococcus and Prochlorococcus groups)are extremely abun-dant in surface waters and contribute a large fraction of pho-tosynthesis and DIN demand (19).Some of these microbes do not contain assimilatory nitrate reductase genes (G.Rocap,rimer,merdin,E.,S.Stilwagon,and S.W.Ch-isholm,Abstr.Aquatic Sci.Meet.,p.120,2001).The high-and low-light-adapted Prochlorococcus spp.(67),divinyl chloro-phyll a -containing cyanobacteria common in oceanic waters (22),differ in their abilities to use N sources (G.Rocap,rimer,merdin,E.,S.Stilwagon,and S.W.Chisholm,Abstr.Aquatic Sci.Meet.,2001).The high-light strain (strain MED4)lacks both nitrate reductase genes and nitrite reduc-tase genes and can grow only on ammonium,whereas the low-light strain (strain MIT9313)lacks only nitrate reductase genes and can grow on nitrite or ammonium (G.Rocap,rimer,merdin,E.,S.Stilwagon,and S.W.Chisholm,Abstr.Aquatic Sci.Meet.,2001).Previously,it was assumed that microorganisms generally could use either nitrate or am-monium and that organisms differed in kinetics of nitrate or ammonium uptake and utilization (29),but now it is clear that some microorganisms may not be able to use nitrate.It is now recognized that bacteria also can play a role in nitrate uptake.It is now known that assimilatory nitrate reduc-tase genes are dispersed among marine bacterial strains as well as among phytoplankton (1).Thus,in complex microbial com-munities in situ,the roles of different microorganisms in up-take of nitrate and ammonium differ due to genetic as well as biochemical and physiological constraints.The major source of nitrate in the ocean surface is diffusion and upwelling of nitrate-rich deep ocean water.In recent years,it has been shown that even this process,which would appear to be driven largely by physical forcing,has an important mi-crobiological component.Many large oceanic diatoms that sometimes form large mats migrate to great depth to obtain nutrients from the nutrient-rich deep water (66),only to return to the surface carrying with them nitrate (99).Migrating Rhi-zosolenia mats may transport an average of 20%,ranging up to 78%,of the upward diffusive flux of nitrate (99).Thus,biolog-ical controls are involved even in the upward movement of nitrate from deep water (Fig.1,pathway D).N LIMITATIONIt is often assumed that N limits the productivity of phyto-plankton in the oceans.This conclusion is based on relatively few studies,most of which assess N limitation indirectly (39).It is quite dif ficult to demonstrate N limitation experimentally.Ideally,one would determine this from a simple bottle exper-iment in which different nutrients are added to each bottle.In ocean studies this can be dif ficult,since microbial communities change during bottle incubations (26).Techniques that directly assay biochemical or physiological targets can provide incuba-tion-independent information on the nutritional status of as-semblages and even individual cells.A number of cellular and molecular markers for nutritional de ficiencies have been de-veloped (36,56,88).Molecular and immunological techniques have provided ways of investigating natural communities,for example,by using probes for the N regulator in cyanobacteria,ntcA (57),or cell surface proteins that are expressed under N limitation (76).Detection of transcripts for ntcA indicated that marine Synechococcus spp.in the Gulf of Aqaba were not nitrogen stressed and were using primarily regenerated nitro-gen.This approach has not yet been applied in open-ocean oligotrophic environments but could be used to determinetheFIG.2.Diagrams of classical and present views of the N cycle in the surface waters of oligotrophic oceans.The composition of the dissolved N pool is shown with approximate relative concentrations of inorganic and organic constituents indicated by the size of the box.Dashed lines indicate transformations and processes included in the newer view of nitrogen cycling.(A)Some phytoplankton use simple organic compounds as a source of nitrogen.(B)There are multiple species of phytoplankton (cyanobacteria)in the open ocean that fix N 2.(C)Bacteria can compete for nitrate and ammonium.(D)Bacteria can excrete urea and can also be a source of high-molecular-weight DON.(E).Some oceanic bacterioplankton appear to fix N 2.V OL .68,2002MINIREVIEW 1017on November 14, 2014 by NANKAI UNIVERSITY/Downloaded fromseasonal and depth variations in nitrogen limitation of cya-nobacteria in the open ocean.HETEROTROPHIC BACTERIAThe primary role of bacteria in the N cycle was presumed to be the release of inorganic N(NH4ϩ)during the decomposi-tion of organic matter,thereby recycling N(and other nutri-ents)to phytoplankton(Fig.1,pathway B;Fig.2).There is now an appreciation for the role of bacteria in the“microbial loop”(3,4),which emphasizes the role of bacteria in trophic trans-fers through a micrograzer(protozoan)food chain.This food chain results in transfer of carbon,N,and other nutrients from dissolved organic matter into the food web.However,this picture is complicated by recentfindings regarding the metab-olism of heterotrophic marine bacteria(for a review,see ref-erence49)and the composition and sources of organic matter (62,63).Bacteria can use DIN as well as organic matter(Fig. 1,pathway A;Fig.2)and thus might even compete with phy-toplankton for inorganic N(50).Whether or not bacteria take up inorganic N probably depends on the C:N ratio of the substrates being used for growth(34).Bacterial regeneration of N during the mineralization of organic matter also depends on the C:N ratio of cell material relative to substrate availabil-ity(49),and thus,whether bacteria provide N or compete with primary producers for N varies substantially in space and time (49).Interestingly,growth rates of bacteria are consistent with the uptake of N from dissolved free amino acids and ammo-nium,indicating that the larger pool of dissolved organic N is not a major source of N for growth(49).Bacteria can take up DIN while simultaneously liberating NH4ϩin decomposition (96).Thus,bacteria in the ocean can be competing for NH4ϩ, regenerating NH4ϩ,or both(Fig.2).It is unclear how both processes are occurring simultaneously,but one explanation is that different members of the microbial assemblage are re-sponsible for different processes(49).DONDissolved organic N(DON)compounds include a wide range of chemical compounds varying in size,complexity,and resilience to degradation.The characterization of dissolved organic matter has long been a challenge to the chemist but perhaps even more so to the microbial ecologist,since it is impossible to trace the metabolism of all of the different com-pounds even if all of the chemical structures and concentra-tions are known.DON can be a large pool in the oceans(3to 7M[12])and an even larger one in coastal waters(90). Important,but usually minor,constituents of DON are amino acids and urea[CO(NH2)2],which are readily used by bacteria and some phytoplankton.Amino acids,either dissolved and free or combined in oligopeptides,are important sources of organic C and N for bacteria(49).Although rates of urea production and catabolism have been measured in marine en-vironments,relatively little is known about the microbiology of urea metabolism in marine systems.Bacteria can be a source or a sink for urea(Fig.2).Urease is a nickel-containing mul-tisubunit metalloprotein encoded by the ure genes(38),which have been characterized in eukaryotes,cyanobacteria,and het-erotrophic bacteria and recently reported in autotrophic nitri-fiers(T.E.Koper,J.M.Norton,and M.G.Klotz,Abstr.9th Int.Symp.Microbial Ecol.,P.08.059,2001).Studies of the diversity of urease genes in the environment are likely to pro-vide interesting information on the distribution of urea utili-zation capabilities in natural assemblages(K.Baker and J. Collier,unpublished data).The definition of“dissolved”organic N has always been a tenuous distinction from that of“particulate”organic nitrogen. This has now been shown to be an artificial distinction,with interactions between organic molecules creating a complete continuum from small dissolved molecules to polymers,col-loids,and particles.Chemical interactions among these mole-cules can result in rearrangements and interactions that form macromolecular gel structures(21).These networked struc-tures,which can form and dissociate biologically or abiologi-cally,can make complex microenvironments that have been suggested to be important for ecological interactions at some microbial scales(3).DON was previously assumed to be primarily high-molecu-lar-weight condensation products,including heterocyclic com-pounds,with a smaller fraction composed of more degradable compounds,including protein,and dissolved free and com-bined amino acids(Fig.2).Recent studies indicate that a significant fraction of DON is composed of amide N(62)and that a fraction originates from bacterial cell walls(63).This finding reverses the perception that phytoplankton are the direct primary source for organic matter through excretion, decomposition,and lysis.Thus,bacteria metabolize organic matter that originates from photosynthetic microorganisms but are now both a source of and a sink for the DON pool(Fig.2). Chemical and photochemical reactions have been found to be important in influencing the quantity and quality of organic matter.Rates of photolysis can equal or exceed bacterial de-composition rates(10).Despite this new perspective on the source and nature of the bulk DOM in the sea,it is abundantly clear that some small fraction of the total DON pool is rapidly cycled by phytoplank-ton and bacteria in the photic zone(Fig.2).A large fraction of the DIN assimilated by phytoplankton can be released as DON within a few hours,probably due to grazing disruption of cells (Fig.2)(9,104).To complicate matters further,there is now evidence that phytoplankton and cyanobacteria can assimilate some small labile components of DON.Thus,the primary producers may not be restricted to exploitation of the DIN pool but may be an important sink for DON as well(Fig.2) (77,78).Release and uptake of DON on a scale of hours or less are probably important for the more labile DON fractions such as protein,peptides,and free amino acids,which are the DON components on which metabolic studies have focused in the past.Bacterioplankton can excrete proteolytic enzymes that digest peptides and proteins so that the monomers or oligo-nucleotides can be taken up by the cell and metabolized(41, 42,91).There are substantial differences in activities and tem-perature responses of these enzymes in different ocean regions, with implications for how and where bacteria are regenerating inorganic N(24).These studies emphasize that the composi-tion of organic matter(e.g.,polysaccharides and sugars versus proteins and amino acids)and the variability among different ocean regions may be important in understanding the role of1018MINIREVIEW A PPL.E NVIRON.M ICROBIOL.on November 14, 2014 by NANKAI UNIVERSITY /Downloaded frombacteria and dissolved organic matter in ocean carbon cycling (24)but also in N regeneration.DON and its metabolism are probably among the most poorly understood parts of the ma-rine N cycle.New methods that combine the use of radiotrac-ers and autoradiography withfluorescence in situ hybridization have begun to provide information on which types of bacteria are metabolizing different organic compounds(75).Given the magnitude of the DON pool and the dependence of hetero-trophic bacteria on dissolved organic matter,this topic de-serves much more attention.Since DON is composed of many different organic substrates,the uptake and metabolism of DON must involve multiple transport proteins and extracellu-lar and intracellular enzymes.The study of DON metabolism is likely to profit from advances in genomic research from the information gained about the diversity of catabolic pathways in cultivated and uncultivated microorganisms.N2FIXATIONBiological N2fixation,the reduction of atmospheric N2to ammonia,is catalyzed by a diverse set of microorganisms.In the marine environment,N2fixation rates are highest in a few specific habitats such as benthic cyanobacterial mats(40).In the oceanic biome,there are relatively few known N2fixers despite the fact that DON concentrations are extremely low in many parts of the ocean.Blooms offilamentous N2-fixing cya-nobacteria often exploit N-limiting conditions in lakes and sometimes estuaries,and so it is a curious paradox that there are so few obvious nitrogen-fixing microorganisms in the ocean (43).An exception is the Baltic Sea,where extensive blooms of filamentous heterocyst-forming cyanobacteria(Aphanizom-enon and Nodularia)frequently occur and have occurred for thousands of years(5).Within the past decade,research on N2fixation in the sea has received increasing attention(47),since the role of the ocean in Cflux is linked to nutrient cycling. In the open ocean,thefilamentous nonheterocystous cya-nobacterium Trichodesmium is common in tropical and sub-tropical waters(13,74).This organism is particularly interest-ing because it forms macroscopic aggregates offilaments,is buoyant due to gas vacuoles,andfixes N2only in the light. Most cyanobacteria segregate O2-sensitive N2fixation from O2 evolved through photosynthesis byfixing N2during the night or in specialized cells,called heterocysts,where photosystem II activity is reduced or absent.Trichodesmium is one of a few known species that evolve O2simultaneously with N2fixation without an obvious mechanism to avoid O2inactivation.Tri-chodesmiumfixes N2only during the day,and this cycle is regulated by the synthesis of nitrogenase under the control of a circadian clock(20).There are various theories and hypoth-eses regarding the mechanisms involved in simultaneous N2fixation and photosynthesis,including the possible division of labor among cells that are morphologically similar(31,45),but the mechanisms whereby Trichodesmiumfixes N2aerobically are still not completely understood(13).The Trichodesmium nitrogenase protein is similar phylogenetically to that of other cyanobacterial diazotrophs and is not likely to be more O2 resistant than other nitrogenases(116).The molecular biology of N2fixation in Trichodesmium has been reviewed elsewhere (116).Perhaps the next most abundant diazotrophs in oceanic wa-ters are the heterocyst-forming cyanobacterial symbionts of diatoms(98).These symbionts have not been successfully maintained in culture for extended periods of time,and so relatively little is known about the biology of the symbiotic interactions between the diatom and cyanobacteria.However, the symbiont-containing diatoms can form large aggregates that can be abundant in oligotrophic waters.These diatoms can form extensive blooms(17)that can be significant sources of N in the mixed layer of the ocean.For years,it was believed that Trichodesmium and the sym-bionts of diatoms were the major N2fixers in the open ocean. However,a number of recent studies have highlighted imbal-ances in N budgets that indicate that higher rates of N2fixation are occurring in the open ocean than was previously estimated (37,59,64).This conclusion is based on biogeochemical cal-culations rather than direct measurements of N2fixation rates or observed distributions of microorganisms but has led to a reevaluation of N2fixation in the sea.Evidence of diverse bacterial and cyanobacterial N2-fixing microorganisms in the Atlantic and Pacific Oceans based on amplification of nitroge-nase genes from bulk water samples has recently been reported (115,118).Nitrogenase genes obtained from the Hawai’i Ocean Time(HOT)series long-term monitoring site at Station ALOHA in the North Pacific Subtropical Gyre were most closely related to nitrogenase genes of unicellular cyanobacte-rial genera that are not typically reported in oceanic waters. The cyanobacterial nifH phylotypes were detected at several times of year and were ultimately shown to be expressing nitrogenase genes(119).Thefinding of the genes led to the microscopic observation of cells that looked similar to the expected morphology(3to8m in diameter,spherical)and subsequent cultivation of N2-fixing isolates(119).Unicellular cyanobacteria with this morphology have been reported from the South Pacific Ocean(71),the Baltic Sea(108),and,pre-viously,from Station ALOHA(11).Estimates of the concen-trations and nitrogenfixation activity of these organisms at Station ALOHA indicate that they could equal or exceed the contribution of Trichodesmium(119),and they have been ob-served elsewhere in the world’s oceans as well.It is not well understood what controls the distribution and activity of diazotrophs in the sea.The distributions of Tri-chodesmium and some other cyanobacteria appear to be cor-related with water temperature(15).It could be that dia-zotrophs are limited by the availability of Fe,a metal that is a component of many proteins in addition to nitrogenase(both the MoFe and Fe components).Fe distributions in the world’s oceans are controlled to a large extent by aeolian transport of dust.Temporal and spatial variation in Fe supply may result in oscillation between N and P limitation of the oceans through its effect on N2fixation(111).However,there does not appear to be a correlation between Fe concentration and Trichodes-mium abundance and activity,at least in the Atlantic Ocean near Bermuda(85).NITRIFICATION/DENITRIFICATION Isolations of nitrifying and denitrifying bacteria using conventional enrichment techniques have provided extensive culture collections on which our understanding of the bio-chemistry and ecology of these processes is based.Chemolitho-V OL.68,2002MINIREVIEW1019on November 14, 2014 by NANKAI UNIVERSITY /Downloaded fromautotrophic nitrification is rather restricted in its occurrence and is represented in culture by25species of ammonia oxidiz-ers in the beta and gamma subdivisions of the Proteobacteria and by eight species of nitrite oxidizers in the alpha,beta,and gamma subdivisions of the Proteobacteria(51).Molecular phy-logeny has supported this generalization,showing that both ammonia-oxidizing bacteria(AOB)and nitrite-oxidizing bac-teria belong to a small number of coherent groups(94). Thefirst AOB to be cultivated from the marine environment was Nitrosocystis oceanus(109),now called Nitrosococcus oce-ani.Strains of N.oceani have been obtained from several locations(14),and it has been detected in seawater by immu-nofluorescence(105)and by PCR amplification(B.B.Ward and G.O’Mullan unpublished data)and in Antarctic lakes by immunofluorescence hybridization andfluorescence in situ hy-bridization(101).Members of the Nitrosomonas/Nitrosospira group appear to dominate most terrestrial and aquatic envi-ronments,including marine sediments(52,73,107,110).Purk-hold et al.(79)surveyed all published16S rRNA and amoA sequences,from both cultures and environmental clones,and concluded that although much diversity among AOB remains to be cultured,it is unlikely that entirely novel species will be discovered,at least in the case of the beta-subdivision AOB. However,this does not preclude the existence of entirely novel AOB that are not detected by probes based on known groups. Nitrite oxidizers have received somewhat less attention,and culture collections are dominated by Nitrobacter strains.Re-cent work in various wastewaters,however,has shown that Nitrospira is the dominant group in these environments(32). For both AOB and nitrite-oxidizing bacteria,phylogeny and functionality appear to be well correlated,making these groups attractive for molecular phylogeny studies despite their slow and fastidious growth habits in culture.Research on nitrifiers, particularly AOB,based on16S rRNA genes and functional genes has proliferated in recent years and was recently re-viewed(53).Denitrifying bacteria are the opposite of nitrifiers in many ways;denitrification ability is found in heterotrophic oppor-tunists and in chemoautotrophs,is widespread among Bacteria and Archaea,and has even been reported in Eukarya(120).16S rRNA and functional gene phylogenies are not congruent for the denitrifiers,implying substantial horizontal gene transfer of the functional genes over time(D.P.Martino and B.B.Ward, Aquatic Sciences Meeting:Limnology and Oceanography:nav-igating into the next century,abstract book,p.117,1999;B. Song and B.B.Ward,submitted for publication).Environmen-tal research on the diversity of denitrifiers has focused,there-fore,on the functional genes involved in the denitrification pathway,mainly nitrite reductase(7,8)and nitrous oxide re-ductase(86,87).Dissimilatory reduction of nitrate to ammonium is often ignored in the marine realm but could be important in sedi-ments in which fermentative bacteria,with whose metabolism it is often associated,are likely to be found.Christensen et al.(23)found it to be a significant nitrate sink only in sediments with very high organic carbon loading,and Bonin et al.(6) suggested that it could be important in coastal sediments,and it is unlikely to occur in the water column.Thus,we refer here mainly to respiratory denitrification,in which nitrate is re-duced sequentially to nitrite,nitric oxide,nitrous oxide,and nitrogen gas.Chemical distributions in marine sediments and the water column indicate that nitrification is an obligately aerobic pro-cess and that denitrification is an obligately anaerobic process. Although the oxygen requirements and tolerances vary among isolates,these requirements are reflected in the physiology of cultivated nitrifiers and denitrifiers,which are predominantly obligate aerobes and facultative anaerobes,respectively.Nitri-fication and denitrification are often coupled across oxic/an-oxic interfaces in both sediments and suboxic waters,leading to the loss offixed nitrogen via mineralization,oxidation,and denitrification(72).Because the deep ocean contains high nitrate concentrations,it was long assumed that nitrification occurred in that environment.However,direct rate measure-ments using15N tracer techniques have consistently shown that most water column nitrification occurs in the lower portion of the euphotic zone and that the nitrate produced there can supply a large fraction of phytoplankton nitrate demand(27, 106).The recent description of aerobic denitrification by novel strains and“conventional”denitrifying bacteria introduces a new link in the N cycle.Aerobic denitrification wasfirst de-scribed in Paracoccus pantotrophus(formerly Thiosphaera pan-totropha[81]),and like others possessing this ability,P.pan-totrophus is also a heterotrophic nitrifier.The nitrite generated by the oxidation of ammonia can be released into the medium or denitrified to N2gas.Denitrification of nitrite or nitrate to N2can occur with atmospheric levels of O2(82).P.pantotro-phus was originally isolated from wastewater;its ability to deni-trify aerobically in batch culture,as well as that of several other conventional heterotrophic denitrifiers,has been confirmed (82).Su et al.(93)reported aerobic denitrification by a strain of Pseudomonas stutzeri at rates and with oxygen tolerance greatly exceeding those reported for P.pantotrophus.The pro-cess may be common in isolates,but its significance in the environment remains uncertain,and even in cultures questions still remain.For example,the enzymology of the aerobic path-way is unknown.While P.pantotrophus expresses the nitrite reductase gene(nirS)under anaerobic conditions,it is not expressed under aerobic conditions(65),and thus,the mech-anism for aerobic reduction of nitrite remains unknown. AOB also perform a subset of the conventional set of deni-trifying reactions,reducing nitrite to NO and N2O.The process occurs aerobically but apparently is enhanced at low oxygen concentrations(35,60).These gases are also intermediates in denitrification but could derive from nitrification under a low or nearly zero oxygen concentration.Both N2O and NO are important in atmospheric processes;they contribute to green-house warming and to catalytic destruction of stratospheric ozone.Thus,understanding which processes are responsible for their production could prove to be important in under-standing or potentially regulating theirfluxes.It appears that chemoautotrophic AOB produce N2O and NO by using a pathway that is essentially identical to the classical denitrification pathway.First,ammonia is oxidized to nitrite and some of the nitrite is reduced to N2O.The reduc-tions are catalyzed by enzymes that are encoded by nitrite reductase and NO reductase genes that are homologous to the nirK and norB genes of conventional denitrifying bacteria(ref-1020MINIREVIEW A PPL.E NVIRON.M ICROBIOL.on November 14, 2014 by NANKAI UNIVERSITY /Downloaded from。
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a rX iv:mat h /511477v1[mat h.GT]21N ov25MILNOR’S ISOTOPY INV ARIANTS AND GENERALIZED LINK HOMOTOPY THOMAS FLEMING 1AND AKIRA YASUHARA Abstract.It has long been known that a Milnor invariant with no repeated index is an invariant of link homotopy.We show that Milnor’s invariants with repeated indices are invariants not only of isotopy,but also of self C k -moves.A self C k -move is a natural generalization of link homotopy based on certain degree k clasper surgeries,which provides a filtration of link homotopy classes.1.Introduction In his landmark 1954paper [8],Milnor introduced his epinonomous higher order linking numbers,and proved that when the multi-index I had no repeated index,the µ(I )were invariants of link homotopy.In a follow up paper [9],Milnor explored some of the properties of invariants with repeated nor’s invariants have been studied extensively since that time.For string links,these invariants are known to be finite type [1,7],and in fact related to (the tree part of)the Kontsevich integral in a natural and beautiful way [4].By work of Habiro [6],the finite type invariants of knots are intimately related to claper surgery.Taking the view that link homotopy is generated by degree one clasper surgery on the link where both leaves of the clasper are on the same component,is it possible that Milnor’s isotopy invariants have some relation with clasper surgery?The answer is yes.Let us define a self C k -move on a link L to be a degree k simple tree clasper surgery on L where all leaves of the clasper are on the same component.If L ′is obtained from L by a sequence of self C k -moves,we call L and L ′self C k -equivalent .These moves were introduced in [15],and as we have mentioned,self C 1-equivalence is link homotopy.For an n -component link,Milnor invariants are specified by a multi-index I ,where the entries of I are chosen from {1,...n }.Let r (I )denote the maximumnumber of times that any index appears.A Milnor invariant µ(I )is called realizable if there exists a link L with µL (I )=0.Our main result is the following.Theorem 2.1Let µ(I )be a realizable Milnor number.Then µ(I )is an invariant of self C k -equivalence if and only if r (I )≤k .Notice that a self C k -equivalence can be realized by self C k ′-moves when k ′<k ,and thus self C k -equivalence classes form a filtration of link homotopy classes.Mov-ing to larger and larger k provides more and more information about the structure of isotopy classes of links.2THOMAS FLEMING1AND AKIRA YASUHARAThe classification of links up to link homotopy has been completed by Habegger and Lin[5].However,very little is known about the structure of links under these higher order moves.Nakanishi and Ohyama have classified two component links up to self C2-equivalence[11,12,13],and it is known that boundary links are self C2-equivalent to the trivial link[16].It is well known that Milnor’s link homotopy invariants vanish if and only if the link is link homotopic to the unlink.However,in Example3.3we will produce a boundary link that is not self C3-equivalent to the unlink.As all Milnor invariants vanish for boundary links,this example demonstrates that self C k-equivalence be-haves very differently for k>1.Much work remains to be done before we have a clear understanding of self C k-equivalence.2.Invariance under self C k-movesWe define a self C k-move to be surgery on a simple,tree-like degree k clasper, all of whose leaves are on the same component.Similarly,L is self C k-equivalent to L′if L′can be obtained from L by a sequence of self C k-moves.Note that link homotopy is the same as self C1-equivalence,and the self-delta equivalence of Shibuya[14]is the same as self C2-equivalence.For an n-component link,Milnor invariants are specified by a multi-index I, where the entries of I are chosen from{1,...n}.Let r(I)denote the maximum number of times that any index appears.A Milnor invariantµ(I)is called realizable if there exists a link L withµL(I)=0.Theorem2.1.Letµ(I)be a realizable Milnor number.Thenµ(I)is an invariant of self C k-equivalence if and only if r(I)≤k.Proof.Suppose that r(I)≤k and L is self C k-equivalent to L′.Since Milnor’s invariants are well known to be isotopy invariants,it suffices to check a single self C k-move realized by a clapser c.The case k=1is that of link homotopy,and was proven by Milnor in[8]. Thus,we need only consider the case k≥2.To calculateµL(I)(resp.µL′(I)) where I has repeated indices,we may instead study the link homotopy invariants of L′),the link obtained by taking the appropriate number of zero framed parallels of the components of L(L′)[9].In particular,if component i ofL (i1,i2...i m)=µL(h(i1),h(i2)...h(i m)).Since k≥2,the self C k-move preserves the framing of the components of L. That is,the clasper surgery carrying L to L′carries L′.The self C k-move is realized by surgery on c,that is,a clasper with k+1leaves where all of these leaves land on the same component of L.Replace L withL has at most k copies each component of L,so each leaf of the clasper c grips at most k components ofL.Use the zip construction on c to produce simple claspers onL,and so is realized by link homotopy.See Proposition3.1.We have shown that L′,soµL′for all link homotopyinvariants.Hence,when r(I)≤k,µL(I)=µL′(I).MILNOR’S ISOTOPY INV ARIANTS AND GENERALIZED LINK HOMOTOPY3 Suppose that r(I)>k.Since the Milnor invariant is realizable,by work of Cochran[2],there exists a link L whereµL(I)=0,and all lower order Milnor invariants vanish.Such a link can be obtained by Bing doubling a Hopf link along a trivalent tree whose leaves are labeled by I,and then band summing components with the same label.This process is the same as a simple tree clasper surgery on the unlink,using the same tree,where leaves labeled i grasp the i th component.Since some index i in I is repeated at least k+1times,this clasper has at least k+1leaves on component i and so can be realized by self C k-moves.See Proposition3.1.As all Milnor invariants of the unlink vanish,we have thatµL(I)is not a self C k invariant when r(I)>k.We end this section with an example that demonstrates the power of our meth-ods.Self C2-equivalence of links is equivalent to self-delta equivalence,a relation that has been extensively studied.Much of this work relies on invariants of self C2-(self-delta)-equivalence that are based on the coefficients of the Conway polynomial.Let L be the Whitehead double of the Borromean rings(equivalently the Bing double of the Whitehead link).See Figure1.The Alexander polynomial of this link is trivial1,butµL(123123)=1.Thus,L has trivial Conway polynomial,but is not self C2-equivalent to the unlink.Figure1.The Whitehead double of the Borromean rings.3.Further resultsIn the proof of Theorem2.1we used the fact that simple clasper surgery with multiple leaves on a single component could be converted to a self C k-equivalence. In this section,we willfind a partial converse to that fact and demonstrate its usefulness in the study of self C k-equivalence.We say L and L′are C k m-equivalent if L can be transformed into L′by ambient isotopy and degree m simple tree clasper surgery,where at least k leaves of the clasper grasp component i.We may now state the fact mentioned above precisely.4THOMAS FLEMING1AND AKIRA YASUHARAProposition3.1.If L is C k+1m-equivalent to L′then L is self C k-equivalent to L′.Proof.We will work by induction.The base case is m=k,and is trivial.Suppose now that c is a clasper representing a C k+1m+1move.We will show that itis C k+1m -equivalent to a C k+1mmove.Begin with the clasper c.Introduce a clasper c′representing a C k+1m -move as in Figure2(2).We choose c′so that by reversing thezip construction,we obtain the clapser with boxes shown in Figure2.By Move12 of[6],we obtain the clasper c′′,which is degree m and has k+1feet on componenti.Thus,we can reduce any C k+1m -move to a sequence of C k+1k-moves,which ismerely a self C k-equivalence.Figure2demonstrates the necessary moves.One mayfind it easier to read this figure from right to left.Figure2.Reducing a C k+1m+1clasper to C k+1mclaspers.The sym-bol s(s−1)represents a positive(negative)half twist.Proposition3.1is a useful,though elementary,application of the properties of claspers.Less obvious,however,is that in certain circumstances we have a converse. Proposition3.2.Let L be an n component Brunnian link,and U the n componentunlink.Then L is self C k-equivalent to U if and only if L is C k+1n+k−1-equivalent toU.Proof.The‘if’part follows from Proposition3.1.We will show the‘only if’part. The idea of this proof is similar to that of Theorem1.2in[10].While in[10]they use the‘band description’defined in[17],we will use clasper theory.Assume that L and U are self C k-equivalent.Let L i be the i th component of L.We can describe L as the unlink O1∪···∪O n with claspers c{i},j representing the self C k-moves on the i th component.Since L is Brunnian,L\L1is the unlink. Thus,L is obtained from the trivial link by C k-moves corresponding to c{1},j and by crossing changes between O1and c{i},j(i≥2).MILNOR’S ISOTOPY INV ARIANTS AND GENERALIZED LINK HOMOTOPY5 Passing a clasper through O1results in two new claspers,one of degree k+1 which has a new leaf on O1,and one of degree k which does not.Call the formerc{1i},j,and the later c′{i},j .See Figure3.We can now express L as the unlink O1∪···∪O n with claspers c{1},j of degreek,claspers c′{i},j (i≥2)of degree k and claspers c{1i},j(i≥2)of degree k+1.Let L′be the link obtained by surgering the unlink O1∪···∪O n only on thec′{i},j .By the construction of the c′{i},j,L′=L\L1 O1.Since L is Brunnian,L′is the unlink,and so the claspers c′{i},j act trivially.Thus we can express L as the unlink O1∪···∪O n with claspers c{1},j of degree k and claspers c{1i},j(i≥2)of degree k+1that have leaves on O1∪O i.Since L\L2is the unlink,repeating the argument above shows that L is expressed as the unlink O1∪···∪O n with claspers c{12},j of degree k+1that have leaves on O1∪O2and claspers c{12i},j(i≥3)of degree k+2that have leaves on O1∪O2∪O i.Repeating these process for each L i,finally,we can express L as the unlink O1∪···∪O n with claspers c{12,...,n},j of degree k+n−1that have leaves on O1∪O2∪···∪O n.Hence L is C k+n−1-equivalent to the unlink.Figure3.Altering the clasper c{i}j to reflect a crossing changebetween that clasper and O1.Figure4illustrates the argument of Proposition3.2for the Whitehead link.The Whitehead link is Brunnian and link homotopic to the unlink.The center image of Figure4shows it as the unlink with one self C1clasper.The right hand image shows the C2-equivalence of the Whitehead link and the unlink.Example 3.3.The link in Figure5is obtained from Whitehead doubling both components of the Hopf link and is a boundary link,so all Milnor invariants vanish. However,examine the Jones polynomial.J(L)=q−92+q−52−q32−2q72We obtained the Jones polynomial from the Knot Atlas2and by calculation using Knot3.Evaluating the third derivative of J(L)at1,we obtain a value distinct from that of the third derivative of Jones polynomial of the unlink at2.Since this is a6THOMAS FLEMING1AND AKIRA YASUHARAFigure4.Proposition3.2for the Whitehead link.finite type invariant of degree three,L is not C4-equivalent to the trivial ing Proposition3.2we see that it is not self C3-equivalent to the trivial link.In fact, since L is Brunnian,if L is self C k-equivalent to a split link,it is self C k-equivalent to the trivial link.Thus,L is not self C3-equivalent to a split link.Figure5.The Hopf link with both components Whitehead doubled.Example3.3is interesting for other reasons.It is well known that Milnor’s link homotopy invariants vanish if and only if L is link homotopic to the unlink. The example above shows that a similar statement is not true for Milnor’s self C3 invariants.While the link in example is a boundary link,it is not self C3-equivalent to the trivial link or even a split link.It would be interesting to know what the vanishing of Milnor’s self C k invariants implies about the self C k-equivalence class of L.For two component links,vanishingµL(12)andµL(1122)implies that the link is self C2-equivalent to the unlink[13].Shibuya and the second author have recently shown that all boundary links are self C2-equivalent to the unlink[16],but whether the vanishing of Milnor’s self C2invariants is sufficient to show that a link is self C2-equivalent to the unlink remains an open question.MILNOR’S ISOTOPY INV ARIANTS AND GENERALIZED LINK HOMOTOPY7References[1]D.Bar Natan,Vassiliev homotopy string link invariants,J.Knot Theory Ramif.4(1995)13-32[2]T.Cochran,Derivatives of links:Milnor’s concordance invariants and Massey’s products,AMS Memoirs,87No.427(1990)[3]J.Conant and P.Teichner Grope cobordism of classical knots Topology43(2004)119-156[4]N.Habegger and G.Masbaum,The Kontsevich integral and Milnor’s invariants,Topology39(2000)1253-1289[5]N.Habegger and X.S.Lin,The classification of links up to link homotopy,J.Am.Math.Soc.3(1990)389-419[6]K.Habiro Claspers andfinite type invariants of links,Geom.and Top.4(2000)1-83[7]X.S.Lin,Power series expansions and invariants of links in:W.Kazez(Ed.)Geomet-ric Topology,Proceedings of the Georgia International Topology Conference1993;AMS/IP Studies in Advanced Mathematics,(1997)[8]nor,Link groups,Ann.Math.59(1954)177-195[9]nor,Isotopy of links in:Algebraic and Geometric Topology,A symposium in honor ofS.Lefshetz,ed.R.H.Fox,Princeton University Press,New Jersey,(1957),280-306[10]H.A.Miyazawa and A.Yasuhara,Classification of n-component Brunnian links up to C nmove,to appear in Topology Appl.[11]Y.Nakanishi,Delta link homotopy for two component links,Topology Appl.121(2002)169-182[12]Y.Nakanishi and Y.Ohyama,Delta link homotopy for two component links II,J.KnotTheory Ramif.11No.3(2002)353-362[13]Y.Nakanishi and Y.Ohyama,Delta link homotopy for two component links III J.Math.Soc.Japan55No.3(2003)641-654[14]T.Shibuya Self∆-equivalence of ribbon links Osaka J.Math.33(1996)751-760[15]T.Shibuya and A.Yasuhara Self C k-move,quasi self C k-move,and the Conway potentialfunction for links J.Knot Theory Ramif.13(2004)877-893[16]T.Shibuya and A.Yasuhara Boundary links are self delta-equivalent to trivial links,preprint[17]K.Taniyama and A.Yasuhara Band description of knots and Vassiliev invariants Math.Proc.Cambridge Phil.Soc.133(2002)325-343University of California San Diego,Department of Mathematics, 9500Gilman Dr.,La Jolla,CA92093-0112,United States E-mail address:tfleming@Tokyo Gakugei University,Department of Mathematics,Koganei-shi,Tokyo184-8501,JapanE-mail address:yasuhara@u-gakugei.ac.jp。