Quantification of photoperiod effects on growth phleum pratense
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Z U O L I W U , A . O . S K J E L V A˚ G and O . H . B A A D S H A U G * Department of Plant and Environmental Sciences, Agricultural University of Norway, PO Box 5003, N-1432 A˚ s, Norway
end
of
experiment.
Tຫໍສະໝຸດ Baidue
plant
responses
to
photoperiod
were
described
by
the
term
PPR
=
, where eciðPPÀPPcÞ
1 + eðci þdi ÞðPPÀPPc Þ
PP = photoperiod in h, PPc = photoperiod of maximum response, c = characteristic coefficient of main response
INTRODUCTION
Photoperiodism is one of the most significant and complex aspects of the interaction between plants and their environment. It is defined as plant responses to daylength, enabling living organisms to adapt to seasonal changes. For instance, at high latitudes autumnal short days signal the induction of bud dormancy and cold hardiness in perennial plant species. Similarly, in desert species dormancy may be induced by long-day conditions, which are accompanied by the unfavourable environment of water stress (Schwabe and Nachmony-Bascombe, 1963).
The southern cultivar ‘Grindstad’ was more sensitive than the northern one ‘Engmo’. The functional relationships
suggest mechanisms for plants’ daylength responses and latitudinal adaptation.
Received: 16 March 2004 Returned for revision: 14 May 2004 Accepted: 16 June 2004 Published electronically: 11 August 2004
Background and Aims Accurate quantifications of plant responses to photoperiod are useful for physiological
ª 2004 Annals of Botany Company
Key words: Daylength, dry matter, functional relationships, leaf area, leaf elongation, modelling, Phleum pratense, photoperiodic effects, quantification.
Most plants are sensitive to photoperiod, not only for generative development but also in many other aspects, such as seed germination, leaf formation rate, leaf blade length and width expansion, dry matter production and its partitioning. Seed germination of rice, an SDP, was promoted by long days (Bhargava, 1975). Long days may result in an enhanced leaf appearance rate in, for example, wheat (Cao and Moss, 1989). However, in very many cases artificial photoperiod extension with low light intensity has been shown to have little or no significant effect on this rate: for example perennial ryegrass (e.g. Gautier et al., 1999), tall fescue (e.g. Skinner and Nelson, 1995), timothy, meadow fescue (e.g. Virkaja¨rvi and Ja¨rvenranta, 2001), and cocksfoot (Østga˚rd and Eagles, 1971). More-or-less the same results have been observed on leaf unfolding in many other crop plants (e.g. Mauchow and Carberry, 1990; Ritchie and NeSmith, 1991; Volk and Bugbee, 1991; Sadras and Villalobos, 1993; McMaster, 1997).
studies, in growth modelling and in other studies of environmental effects. The objective of the current work was a
mathematical description of photoperiodic influence on plant morphological traits, using functions with few and
growth as expressed by several plant characteristics, such as leaf area development, top and root DM production, as
well as cultivar differences. Green leaf area was more sensitive to photoperiod than above-ground DM production.
Annals of Botany 94: 535–543, 2004 doi:10.1093/aob/mch170, available online at www.aob.oupjournals.org
Quantification of Photoperiodic Effects on Growth of Phleum pratense
interval, d = sensitivity coefficient characterizing course of function beyond the main response interval. PPR was
tested on experimental data for different growth characteristics (i), e.g. size of individual leaves (Yi), identified by their sequential numbers on the main tiller (LN) using the function: Yi = Ybi + aiLN + biLNai (PRR). The growth course was described by the same function, replacing LN with day number of treatment exposure. Key Results and Conclusions The functions described with high precision (r2 > 0Á97) the effect of photoperiod on
The classification of plants according to their photoperiodic responses is usually based on flowering. The two main photoperiodic response categories are short-day plants (SDPs) and long-day plants (LDPs), in which flowering occurs in short or long days, respectively. Both types may respond qualitatively or quantitatively to daylength. Besides these types and day-neutral plants, a few plant species have more specialized daylength requirements; i.e. intermediate-day plants (IDPs), in which flowering occurs only between narrow daylength limits (e.g. 12–14 h for one cultivar of sugarcane) and ambiphotoperiodic species (APPSs) in which flowering occurs only in long
common parameters related to key plant characteristics and typical response patterns.
Methods Two latitudinal cultivars of timothy (Phleum pratense) were studied in a climate chamber experiment at
* For correspondence. E-mail ole.baadshaug@ipm.nlh.no
or short days (e.g. Madia elegans) but not at intermediate daylengths (cf. Taiz and Zeiger, 1998). Most temperate grasses and sedges (Carex spp.) may have a dual daylength requirement for flowering (Heide, 1987, 1994, 1997), for example the plants must first be exposed to short days and thereafter to long days (SD–LD), or the dual requirement is reversed (LD–SD plants, e.g. Bryophyllum daigremontianum; Zeevaart, 1969).
9, 12, 15, 18, 21 and 24 h photoperiods. Seedling growth was recorded by measurements of main tiller leaf tip heights
every other day from the 5–6 leaf stage onwards, and as plant size and dry weight at days 37, 46, 62 and 70, i.e. at the
end
of
experiment.
Tຫໍສະໝຸດ Baidue
plant
responses
to
photoperiod
were
described
by
the
term
PPR
=
, where eciðPPÀPPcÞ
1 + eðci þdi ÞðPPÀPPc Þ
PP = photoperiod in h, PPc = photoperiod of maximum response, c = characteristic coefficient of main response
INTRODUCTION
Photoperiodism is one of the most significant and complex aspects of the interaction between plants and their environment. It is defined as plant responses to daylength, enabling living organisms to adapt to seasonal changes. For instance, at high latitudes autumnal short days signal the induction of bud dormancy and cold hardiness in perennial plant species. Similarly, in desert species dormancy may be induced by long-day conditions, which are accompanied by the unfavourable environment of water stress (Schwabe and Nachmony-Bascombe, 1963).
The southern cultivar ‘Grindstad’ was more sensitive than the northern one ‘Engmo’. The functional relationships
suggest mechanisms for plants’ daylength responses and latitudinal adaptation.
Received: 16 March 2004 Returned for revision: 14 May 2004 Accepted: 16 June 2004 Published electronically: 11 August 2004
Background and Aims Accurate quantifications of plant responses to photoperiod are useful for physiological
ª 2004 Annals of Botany Company
Key words: Daylength, dry matter, functional relationships, leaf area, leaf elongation, modelling, Phleum pratense, photoperiodic effects, quantification.
Most plants are sensitive to photoperiod, not only for generative development but also in many other aspects, such as seed germination, leaf formation rate, leaf blade length and width expansion, dry matter production and its partitioning. Seed germination of rice, an SDP, was promoted by long days (Bhargava, 1975). Long days may result in an enhanced leaf appearance rate in, for example, wheat (Cao and Moss, 1989). However, in very many cases artificial photoperiod extension with low light intensity has been shown to have little or no significant effect on this rate: for example perennial ryegrass (e.g. Gautier et al., 1999), tall fescue (e.g. Skinner and Nelson, 1995), timothy, meadow fescue (e.g. Virkaja¨rvi and Ja¨rvenranta, 2001), and cocksfoot (Østga˚rd and Eagles, 1971). More-or-less the same results have been observed on leaf unfolding in many other crop plants (e.g. Mauchow and Carberry, 1990; Ritchie and NeSmith, 1991; Volk and Bugbee, 1991; Sadras and Villalobos, 1993; McMaster, 1997).
studies, in growth modelling and in other studies of environmental effects. The objective of the current work was a
mathematical description of photoperiodic influence on plant morphological traits, using functions with few and
growth as expressed by several plant characteristics, such as leaf area development, top and root DM production, as
well as cultivar differences. Green leaf area was more sensitive to photoperiod than above-ground DM production.
Annals of Botany 94: 535–543, 2004 doi:10.1093/aob/mch170, available online at www.aob.oupjournals.org
Quantification of Photoperiodic Effects on Growth of Phleum pratense
interval, d = sensitivity coefficient characterizing course of function beyond the main response interval. PPR was
tested on experimental data for different growth characteristics (i), e.g. size of individual leaves (Yi), identified by their sequential numbers on the main tiller (LN) using the function: Yi = Ybi + aiLN + biLNai (PRR). The growth course was described by the same function, replacing LN with day number of treatment exposure. Key Results and Conclusions The functions described with high precision (r2 > 0Á97) the effect of photoperiod on
The classification of plants according to their photoperiodic responses is usually based on flowering. The two main photoperiodic response categories are short-day plants (SDPs) and long-day plants (LDPs), in which flowering occurs in short or long days, respectively. Both types may respond qualitatively or quantitatively to daylength. Besides these types and day-neutral plants, a few plant species have more specialized daylength requirements; i.e. intermediate-day plants (IDPs), in which flowering occurs only between narrow daylength limits (e.g. 12–14 h for one cultivar of sugarcane) and ambiphotoperiodic species (APPSs) in which flowering occurs only in long
common parameters related to key plant characteristics and typical response patterns.
Methods Two latitudinal cultivars of timothy (Phleum pratense) were studied in a climate chamber experiment at
* For correspondence. E-mail ole.baadshaug@ipm.nlh.no
or short days (e.g. Madia elegans) but not at intermediate daylengths (cf. Taiz and Zeiger, 1998). Most temperate grasses and sedges (Carex spp.) may have a dual daylength requirement for flowering (Heide, 1987, 1994, 1997), for example the plants must first be exposed to short days and thereafter to long days (SD–LD), or the dual requirement is reversed (LD–SD plants, e.g. Bryophyllum daigremontianum; Zeevaart, 1969).
9, 12, 15, 18, 21 and 24 h photoperiods. Seedling growth was recorded by measurements of main tiller leaf tip heights
every other day from the 5–6 leaf stage onwards, and as plant size and dry weight at days 37, 46, 62 and 70, i.e. at the