Phytochrome photoreceptors mediate plasticity to light quality in flowers of the Brassicaceae

The family of phytochrome photoreceptors mediates stem-elongation responses to ambient ratios of red?:?far-red light (R?:?FR). Although phytochrome genes are expressed in flowers in addition to vegetative parts, nothing is known about floral plasticity to R?:?FR or the pleiotropic effects of phytochrome genes on flowers. Here, the following floral morphologies were compared: (1) wild-type Arabidopsis thaliana and Brassica rapa plants experiencing high R?:?FR characteristic of sunlight vs. low R?:?FR typical of foliar shade and (2) wild-type and phytochrome-deficient A. thaliana plants. Wild-type A. thaliana exposed to low R?:?FR had diminished petal and pistil lengths but longer filaments for a given petal size than plants experiencing high R?:?FR. Brassica rapa plants had qualitatively similar responses. In comparison to wild-type A. thaliana, mutants lacking phytochrome A had smaller flowers (smaller petals, pistils, and filaments), whereas phytochrome B-deficient mutants exhibited longer filament lengths. These results provide the first evidence that R?:?FR and phytochromes affect floral phenotypes in addition to vegetative ones. Although the ecological relevance remains to be established, the observed plasticity of flowers to R?:?FR may be relevant to individual fitness in some species because stigma and filament positions can affect pollen removal and levels of self-pollination.     

The evolvability of herkogamy: Quantifying the evolutionary potential of a composite trait

Accurate estimates of trait evolvabilities are central to predicting the short‐term evolutionary potential of populations, and hence their ability to adapt to changing environments. We quantify and evaluate the evolvability of herkogamy, the spatial separation of male and female structures in flowers, a key floral trait associated with variation in mating systems. We compiled genetic‐variance estimates for herkogamy and related floral traits, computed evolvabilities, and compared these among trait groups and among species differing in their mating systems. When measured in percentage of its own size, the median evolvability of herkogamy was an order of magnitude greater than the evolvability of other floral size measurements, and was generally not strongly constrained by genetic covariance between its components (pistil and stamen lengths). Median evolvabilities were similar across mating systems, with only a tendency toward reduction in highly selfing taxa. We conclude that herkogamy has the potential to evolve rapidly in response to changing environments. This suggests that the extensive variation in herkogamy commonly observed among closely related populations and species may result from rapid adaptive tracking of fitness optima determined by variation in pollinator communities or other selective factors.     

Quantitative trait loci for floral morphology in Arabidopsis thaliana

A central question in biology is how genes control the expression of quantitative variation. We used statistical methods to estimate genetic variation in eight Arabidopsis thaliana floral characters (fresh flower mass, petal length, petal width, sepal length, sepal width, long stamen length, short stamen length, and pistil length) in a cosmopolitan sample of 15 ecotypes. In addition, we used genome-wide quantitative trait locus (QTL) mapping to evaluate the genetic basis of variation in these same traits in the Landsberg erecta x Columbia recombinant inbred line population. There was significant genetic variation for all traits in both the sample of naturally occurring ecotypes and in the Ler x Col recombinant inbred line population. In addition, broad-sense genetic correlations among the traits were positive and high. A composite interval mapping (CIM) analysis detected 18 significant QTL affecting at least one floral character. Eleven QTL were associated with several floral traits, supporting either pleiotropy or tight linkage as major determinants of flower morphological integration. We propose several candidate genes that may underlie these QTL on the basis of positional information and functional arguments. Genome-wide QTL mapping is a promising tool for the discovery of candidate genes controlling morphological development, the detection of novel phenotypic effects for known genes, and in generating a more complete understanding of the genetic basis of floral development.     

Responses of carbon acquisition traits to irradiance and light quality in Mercurialis annua (Euphorbiaceae): evidence for weak integration of plastic responses

It is often suggested that traits will be integrated, either because of pleiotropy or because natural selection may favor suites of integrated traits. Plant responses to different environments can provide evidence of such integration. We grew Mercurialis annua plants in high-density stands in high irradiance, in neutral shade, and in high red to far-red (R:FR) shade, resulting in environments of high irradiance, low R:FR; low irradiance, low R:FR; and low irradiance, high R:FR. We measured gas exchange, leaf morphology, stem elongation, and biomass traits and tested the prediction that traits within each functional group would show higher trait integration, as evidenced by high correlations among traits within environments, higher correlations of trait plasticity, and lower plasticity of trait correlations. Overall, we found evidence of only moderate integration for some groups of traits. Functionally related groups of traits, or pairs of traits, could be strongly integrated by one criterion but weakly integrated by another of the criteria. Stem elongation traits, though often observed to be strongly integrated in other taxa, showed little evidence of integration. Internode traits exhibited a novel pattern of responses to low R:FR, with increased elongation of the hypocotyl, decreased elongation of the first internode, and no change in the second internode. We propose that these responses to light are more likely to be the result of natural selection than the consequence of constraints imposed by pleiotropy.     

木槿与野西瓜苗花特征和繁育系统的比较研究

为比较分析柱头裂片运动的野西瓜苗和不运动的木槿在花特征和繁育系统上的差异,对其花性状、花粉/胚珠比和不同授粉处理的座果率进行了测定。结果表明:(1)木槿与野西瓜苗在花冠大小、花瓣长及底宽、花萼长与宽、雄蕊长、最上轮雄蕊高度、雄蕊柱长及其底径、花柱长度和雌雄异位间均存在十分显著的差异(P<0.01),但二者的最下轮雄蕊高度间无显著性差异(P>0.05)。(2)木槿的花粉/胚珠比为874.90±20.79,繁育系统属兼性异交类型;野西瓜苗的花粉/胚珠比为24.72±0.68,繁育系统属专性自交类型。(3)木槿自然套袋的座果率为0%,人工自交为2.04%,人工杂交为35.85%;野西瓜苗自动自交和人工自交的座果率均为100%,人工杂交为95.30%。木槿与野西瓜苗在花性状、花粉/胚珠比及花部行为等方面形成了与其繁育系统(兼性异交vs.专性自交)相适应的花特征。     

Modification of flower architecture during early stages in the evolution of self-fertilization

Background and Aims

The evolution of selfing from outcrossing is characterized by a series of morphological changes to flowers culminating in the selfing syndrome. However, which morphological traits initiate increased self-pollination and which are accumulated after self-fertilization establishes is poorly understood. Because the expression of floral traits may depend on the conditions experienced by an individual during flower development, investigation of changes in mating system should also account for environmental and developmental factors. Here, early stages in the evolution of self-pollination are investigated by comparing floral traits among Brazilian populations of Eichhornia paniculata (Pontederiaceae), an annual aquatic that displays variation in selfing rates associated with the breakdown of tristyly to semi-homostyly.

Methods

Thirty-one Brazilian populations under uniform glasshouse conditions were compared to investigate genetic and environmental influences on flower size and stigma–anther separation (herkogamy), two traits that commonly vary in association with transitions to selfing. Within-plant variation in herkogamy was also examined and plants grown under contrasting environmental conditions were compared to examine to what extent this trait exhibits phenotypic plasticity.

Key Results

In E. paniculata a reduction in herkogamy is the principal modification initiating the evolution of selfing. Significantly, reduced herkogamy was restricted to the mid-styled morph and occurred independently of flower size. Significant genetic variation for herkogamy was detected among populations and families, including genotypes exhibiting developmental instability of stamen position with bimodal distributions of herkogamy values. Cloned genets exposed to contrasting growth conditions demonstrated environmental control of herkogamy and genotypic differences in plasticity of this trait.

Conclusions

The ability to modify herkogamy independently of other floral traits, genetic variation in the environmental sensitivity of herkogamy, and the production of modified and unmodified flowers within some individuals, reveal the potential for dynamic control of the mating system in a species that commonly confronts heterogeneous aquatic environments.Key words: Eichhornia paniculata, expressivity, flower morphology, herkogamy, phenotypic plasticity, pleiotropy, population variation, self-fertilization, stigma–anther separation, outcrossing, tristyly     

土壤养分对蝴蝶花的花形态可塑性的影响

为了研究土壤养分对蝴蝶花的花形态可塑性的影响,在田间实验中对蝴蝶花进行了高、中、低养分（N、P、K）水平处理。实验表明,随着土壤养分水平的增加,外轮花瓣长和宽、内轮花瓣长和宽、雌蕊长、雄蕊长、花冠直径均逐渐增加。花各组成部分的形态大小间相关性显著（P=0．05）。结合花部形态特征对环境异质性的生态适应意义,对实验结果进行了讨论。     

Evolutionary analysis of a key floral trait in aquilegia canadensis (ranunculaceae): genetic variation in herkogamy and its effect on the mating system

The mating system of flowering plant populations evolves through selection on genetically based phenotypic variation in floral traits. The physical separation of anthers and stigmas within flowers (herkogamy) is expected to be an important target of selection to limit self-fertilization. We investigated the pattern of phenotypic and genetic variation in herkogamy and its effect of self-fertilization in a broad sample of natural populations of Aquilegia canadensis, a species that is highly selfing despite strong inbreeding depression. Within natural populations, plants exhibit substantial phenotypic variation in herkogamy caused primarily by variation in pistil length rather than stamen length. Compared to other floral traits, herkogamy is much more variable and a greater proportion of variation is distributed among rather than within individuals. We tested for a genetic component of this marked phenotypic variation by growing naturally pollinated seed families from five populations in a common greenhouse environment. For three populations, we detected a significant variation in herkogamy among families, and a positive regression between parental herkogamy measured in the field and progeny herkogamy in the greenhouse, suggesting that there is often genetic variation in herkogamy within natural populations. We estimated levels of self-fertilization for groups of flowers that differed in herkogamy and show that, as expected, herkogamy was associated with reduced selfing in 13 of 19 populations. In six of these populations, we performed floral emasculations to show that this decrease in selfing is due to decreased autogamy (within-flower selfing), the mode of selfing that herkogamy should most directly influence. Taken together, these results suggest that increased herkogamy should be selected to reduce the production of low-quality selfed seed. The combination of high selfing and substantial genetic variation for herkogamy in A. canadensis is enigmatic, and reconciling this observation will require a more integrated analysis of how herkogamy influences not only self-fertilization, but also patterns of outcross pollen import and export.     

Covariance of floral and vegetative traits in four species of Ranunculaceae: a comparison between specialized and generalized pollination systems

Theory predicts that tighter correlation between floral traits and weaker relationship between floral and vegetative traits more likely occur in specialized flowers than generalized flowers,favoring by precise fit with pollinators.However,traits and trait correlations frequently vary under different environments.Through detecting spatiotemporal variation in phenotypic traits (floral organ size and vegetative size) and trait correlations in four Ranunculaceae species,we examined four predictions.Overall,our results supported these predictions to a certain degree.The mean coefficient of variation (CV) of floral traits in two specialized species (Delphinium kamaonense and Aconitum gymnandrum) was marginally significantly lower than that of another two generalized species (Trollius ranunculoides and Anemone obtusiloba).The two specialized species also showed marginally significantly smaller CV in floral traits than vegetative size across the two species.The absolute mean correlation between floral and vegetative traits,or that between floral traits in species with specialized flowers was not significantly lower,or higher than that in generalized plants,weakly supporting the predictions.Furthermore,we documented a large variation in trait correlations of four species among different seasons and populations.Study of covariance of floral and vegetative traits will benefit from the contrast of results obtained from generalized and specialized pollination systems.     

Floral trait variation in fourCyclamen (Primulaceae) species

The purpose of this study is to examine floral trait variation between fourCyclamen species that show variation in their ability to reproduce in the absence of pollinators and in levels of inbreeding. Pollen and ovule production, pollen/ovule ratio, pollen volume, petal length and width, diameter of the corolla mouth, pistil and stamen length, and stigma-anther separation varied significantly between the four study species. Flower, pollen and ovule production, pollen volume and corolla size were generally highest inC. hederifolium, the species with the lowest level of inbreeding (mean F_is = 0.329), intermediate in both species with relatively high levels of pollinator-mediated inbreeding,C. repandum (mean F_is = 0.658) andC. creticum (mean F_is = 0.748), and lowest in the highly inbredC. balearicum (mean F_is = 0.930). The two species with the most different inbreeding coefficients,C. hederifolium andC. balearicum, had lower pollen-ovule ratios and shorter longevities of stigma receptivity and pollen viability thanC. creticum andC. repandum. These patterns of variation in floral traits are discussed in relation to the relatedness, pollination ecology and levels of inbreeding of the four species.     

Morph-specific variation of floral traits associated with reciprocal herkogamy in natural populations of Primula vulgaris and Primula veris

Documenting the morph-specific variation of floral traits associated with reciprocal herkogamy is of special importance for revealing the functional significance of traits in the evolution and maintenance of the heterostylous syndrome. In order to describe the extent and specificity of variation, stigma height, anther height, stigma-anther separation and corolla tube length were measured on 800 flowers collected in two natural populations of Primula vulgaris and P. veris. Beside the almost complete separation of stigma heights between the two morphs, we found appreciable intermorph overlap in anther height and relatively broad range of stigma-anther separation so far reported for heterostylous species. The corolla tube length–stigma-anther separation relationship showed striking difference between the two floral morphs, which supports the hypothesis that length of the corolla tube plays a more important role in positioning the sex-organs in the appropriate distance in the short-styled than in the long-styled morph.     

GENETIC CONSTRAINTS ON THE INDEPENDENT EVOLUTION OF MALE AND FEMALE REPRODUCTIVE CHARACTERS IN THE TRISTYLOUS PLANT LYTHRUM SALICARIA

Here we test whether the potential exists for the independent evolution of allocation to male, female, and attractive functions within a flower. We employed half-sib and parent-offspring regression methods in the tristylous plant Lythrum salicaria to determine whether there is additive genetic variation for characters important to male and female reproductive success and whether genetic correlations could constrain the independent evolution of male and female function. Although significance levels were not consistent among morph types or between populations, there were significant narrow-sense heritabilities for several traits including stamen mass, pistil mass, perianth mass, petal length, and calyx length. Traits that might be under strong stabilizing selection to promote specific pollen transfer, such as stamen and style lengths, had little heritable variation. In the majority of cases in which heritable variation was present, there were positive genetic correlations among floral traits. A strong positive genetic correlation appeared between stamen and pistil mass in the short-styled morph from one of the populations studied. This suggests that selection might not be able to act independently on biomass allocation to male and female flower parts. No evidence of negative genetic correlations appeared that would suggest trade-offs and that could augment a selection response towards sexual specialization. The observed positive correlations could be explained if we consider the “functional architecture” that underlies the covariance structure. If there is more covariance generated by pleiotropic loci controlling overall flower size than at loci controlling male versus female allocation, it could result in the observed positive covariance. At the phenotypic level, we did find significant negative partial correlations between male and female traits when flower size was controlled, but these trade-offs were among rather than within morphs.     

Mapping QTLs influencing rice floral morphology using recombinant inbred lines derived from a cross between <Emphasis Type="Italic">Oryza sativa</Emphasis> L. and <Emphasis Type="Italic">Oryza rufipogon</Emphasis> Griff

To understand the genetic basis of floral traits associated with the mating system in rice, we analyzed pistil, stamen and glume traits using a recombinant inbred line population, derived from a cross between an Asian cultivated rice ( Oryza sativa L.), Pei-kuh, and a wild rice ( Oryza rufipogon Griff.), W1944. Quantitative trait loci (QTLs) affecting floral morphology were detected by composite interval mapping using a linkage map constructed using 147 markers, mostly RFLPs. A total of 7, 4, 14 and 6 QTLs were detected for traits related to pistil, stamen, and size and shape of the glume, respectively. Comparison of 31 QTLs affecting these organs revealed ten QTLs affecting the different organs in four adjacent regions on chromosomes 2, 4, 5 and 10, but most QTLs (68%) were located separately on the whole chromosomes. Although four QTLs for stigma breadth, anther length and thickness of lemma and palea explained more than 25% of the total phenotypic variance, most QTLs (87%) had smaller effects. These results suggest that quantitative variation observed for pistil, stamen and glume traits is controlled by several distinct genes with small effects.     

Physiological responses of spring rapeseed (Brassica napus) to red/far‐red ratios and irradiance during pre‐ and post‐flowering stages

Early shade signals promote the shade avoidance syndrome (SAS) which causes, among others, petiole and shoot elongation and upward leaf position. In spite of its relevance, these photomorphogenic responses have not been deeply studied in rapeseed (Brassica napus). In contrast to other crops like maize and wheat, rapeseed has a complex developmental phenotypic pattern as it evolves from an initial rosette to the main stem elongation and an indeterminate growth of floral raceme. In this work, we analyzed (1) morphological and physiological responses at individual level due to low red/far‐red (R/FR) ratio during plant development, and (2) changes in biomass allocation, grain yield and composition at crop level in response to high R/FR ratio and low irradiance in two modern spring rapeseed genotypes. We carried out pot and field experiments modifying R/FR ratios and irradiance at vegetative or reproductive stages. In pot experiments, low R/FR ratio increased the petiole and lamina length, upward leaf position and also accelerated leaf senescence. Furthermore, low R/FR ratio reduced main floral raceme and increased floral branching with higher remobilization of soluble carbohydrates from the stems. In field experiments, low irradiance during post‐flowering reduced grain yield, harvest index and grain oil content, and high R/FR ratio reaching the crop partially alleviated such effects. We conclude that photomorphogenic signals are integrated early during the vegetative growth, and irradiance has stronger effects than R/FR signals at rapeseed crop level.     

基于同质园栽培的21种淫羊藿属植物花部特征的分类学意义

以采自中国多地并同质园栽培于吉首大学种质资源圃的21种淫羊藿为研究材料,通过观测这些物种的花梗长、花直径、内萼片长宽、蜜距长、雌雄蕊长、花药长、花柱长、胚珠数、花粉大小及花粉量等16个花部特征数量性状和花序类型、有无蜜距、花瓣颜色、内萼片颜色、内萼片与花瓣相对长度及花粉颜色6个质量性状,并对其分类学意义进行探讨。结果表明,(1)21种淫羊藿内轮萼片花瓣状,4枚花瓣除无距淫羊藿外均特化成蜜距,但花部形态种间存在显著差异,尤其体现于花直径、内萼片长宽、蜜距长、雌蕊长、胚珠数、花粉量、花粉直径等特征。(2)主成分分析显示花直径、内萼片长、花药长、胚珠数和花粉直径等性状信息负荷量较大,在该属分类中起主要作用。(3)根据花部特征聚类分析,物种间区分良好,21物种可分为4支:第1支花小,花瓣短于内轮萼片,囊状距或兜状距;第2支花较大,花瓣远长于内轮萼片,长距状;第3支花小,花瓣没有特化形成蜜距;第4支花较小,花丝伸长明显,可达10 mm,花蜜距呈钻状。(4)依据所观察到的花部形态特征编写了淫羊藿属21种植物花部特征分种检索表。研究认为,基于同质园栽培下的淫羊藿属物种的花部特征具有分类学意义,可以为该属分类提供一定依据。     

异质生境对二型花柱植物雪地报春花性状分化的影响

该研究以生长在新疆北部的二型花柱植物雪地报春(Primula nivalis)为研究材料,对7个低海拔(1 657~2 013m)森林居群和7个高海拔(2 400~2 700m)草原居群的花型比例进行调查,对花部特征进行检测,并对长、短花柱植株在不同海拔居群的花性状分化进行了比较研究,以揭示该类群植物的演化与环境的关系。结果表明:(1)在高海拔草原居群中长花柱植株比例高于短花柱植株。(2)比较草原和森林生态系统中的雪地报春居群,长花柱花形态特征间存在显著差异,但短花柱花形态特征间的差异不显著。(3)高海拔草原居群中长花柱花的花冠筒长度、雄蕊长度、雌雄蕊距离、雌雄异位程度以及雌雄蕊空间位置是导致花性状分化的主要因素。(4)在草原居群中,长花柱花的雄蕊、雌雄异位程度及花冠口至雄蕊距离的变化减少了长短花柱花间的互补式对应性。研究认为,在异质生境中长花柱花性状的变化是导致该物种长短花柱花性状分化的主要因素。     

Adaptive Significance of Floral Movement

Since Darwin observed the reconfiguration of pollinia in orchards and referred to it as a function to reduce self-pollination, diverse floral movements have been investigated and various hypotheses have been proposed to explain their adaptive significance. However, adaptive significance of floral movement in some species has yet to be fully explained. Increasing evidence suggests that some floral movements, which have previously been considered as a mechanism to avoid self-pollination, may act as a mechanism to reduce intrafloral male-female interference. In this review, we first explore insect-induced floral movement—such as secondary pollen presentation—that enhances the efficiency and accuracy of pollination. Secondly, we outline the active movements of different floral structures of pistil (style), stamen (filament, anther, pollen), and corolla, such as flexistyly, pollen sliding and catapulting, and anther rotation. Thirdly, we introduce movement herkogamy, which makes herkogamy decrease or increase via the movements of floral structures, especially decreased approach herkogamy via style curvature in the Malvaceae with a monadelphous column. Fourthly, we highlight the different adaptive significances in floral movements for understanding their evolution, including reduction in intrafloral male–female interference, promotion of outcrossing and/or avoidance of self-pollination, delayed autonomous selfing, and tolerance to harsh environments. In particular, we explore the possibility of three functions of one floral movement in one species, which differs from the generally recognized and conventional notion of one floral movement in one species with only one or two functional mechanisms. Finally, we provide perspectives on the challenges and opportunities for using demographic and molecular genetic approaches to (i) study the relative importance and evolutionary mechanism of different adaptive significances in one floral movement; and (ii) simultaneously investigate the floral movement and correlative traits of broader species in related or unrelated families to test how they evolved and the evolutionary relationship between their functions.     

黄瓜去顶苗直接成花的形态学研究

本文对黄瓜0—7天幼苗及去顶后0—8天诱导花芽分化苗与诱导营养芽分化苗的子叶节进行了系统石蜡切片观察,未发现0—7天幼苗的子叶叶腋存在潜伏芽。去顶后1—2天在子叶叶柄基部与切口之间的表皮下细胞分裂形成突起,去顶后6天诱花苗与诱芽苗的突起表现出形态差异,诱花苗突起的上端变钝,而诱芽苗突起的上端成尖锥状。去顶后8天诱花苗在子叶叶柄与切口之间形成完整的花芽。另发现有少量花芽起源于切口处细胞。对去顶后0—6天诱花苗与诱芽苗的子叶节还进行了电镜扫描观察,观察结果与石蜡切片基本一致。     

Minor quantitative trait loci underlie floral traits associated with mating system divergence in Mimulus

The genetic basis of species differences provides insight into the mode and tempo of phenotypic divergence. We investigate the genetic basis of floral differences between two closely related plant taxa with highly divergent mating systems, Mimulus guttatus (large-flowered outcrosser) and M. nasutus (small-flowered selfer). We had previously constructed a framework genetic linkage map of the hybrid genome containing 174 markers spanning approximately 1800 cM on 14 linkage groups. In this study, we analyze the genetics of 16 floral, reproductive, and vegetative characters measured in a large segregating M. nasutus x M. guttatus F2 population (N = 526) and in replicates of the parental lines and F1 hybrids. Phenotypic analyses reveal strong genetic correlations among floral traits and epistatic breakdown of male and female fertility traits in the F2 hybrids. We use multitrait composite interval mapping to jointly locate and characterize quantitative trait loci (QTLs) underlying interspecific differences in seven floral traits. We identified 24 floral QTLs, most of which affected multiple traits. The large number of QTLs affecting each trait (mean = 13, range = 11-15) indicates a strikingly polygenic basis for floral divergence in this system. In general, QTL effects are small relative to both interspecific differences and environmental variation within genotypes, ruling out QTLs of major effect as contributors to floral divergence between M. guttatus and M. nasutus. QTLs show no pattern of directional dominance. Floral characters associated with pollinator attraction (corolla width) and self-pollen deposition (stigma-anther distance) share several pleiotropic or linked QTLs, but unshared QTLs may have allowed selfing to evolve independently from flower size. We discuss the polygenic nature of divergence between M. nasutus and M. guttatus in light of theoretical work on the evolution of selfing, genetics of adaptation, and maintenance of variation within populations.     

Is photomorphogenic shade avoidance adaptive? Perspectives from population biology

Photomorphogenic shade avoidance responses provide an ideal model system for integrating genetic, physiological and population biology approaches to the study of adaptive plasticity. The adaptive plasticity hypothesis predicts that shade avoidance phenotypes induced by low ratios of red to far-red light (R:FR) will have high relative fitness in dense stands, but will suffer a fitness disadvantage at low density. Experiments with transgenic and mutant plants in which photomorphogenic genes are disabled, as well as phenotype manipulation by means of altered R:FR, strongly support the shade avoidance hypothesis. The observation of photomorphogenic ecotypes in different selective environments also suggests that the shade avoidance response has undergone adaptive evolution. Quantitative genetic variation in R:FR sensitivity has been detected in wild populations, indicating that the evolutionary potential exists for response to natural selection. However, evolutionary response may be constrained by genetic correlations among developmentally linked traits. Therefore it cannot be assumed that an observed suite of photomorphogenic responses represents an adaptive optimum for every trait.