Testing the adaptive plasticity of Iris pumila leaf traits to natural light conditions using phenotypic selection analysis

A multivariate selection analysis has been used to test the adaptiveness of several Iris pumila leaf traits that display plasticity to natural light conditions. Siblings of a synthetic population comprising 31 families of two populations from contrasting light habitats were grown at an open dune site and in the understory of a Pinus nigra stand in order to score variation in phenotypic expression of six leaf traits: number of senescent leaves, number of live leaves, leaf length, leaf width, leaf angle, and specific leaf area. The ambient light conditions affected the values of all traits studied except for specific leaf area. In accordance to ecophysiological expectations for an adaptive response to light, both leaf length and width were significantly greater while the angle between sequential leaves was significantly smaller in the woodland understory than at the exposed dune site. The relationship between leaf traits and vegetative fitness (total leaf area) differed across light habitats as predicted by functional hypotheses. The standardized linear selection gradient (β′) for leaf length and width were positive in sign in both environments, but their magnitude for leaf length was higher in the shade than under full sunlight. Since plasticity of leaf length in the woodland shade has been recognized as adaptive, fitness cost of producing plastic change in leaf length was assessed. In both of the available methods used, the two-step and the multivariate regression procedures, a rather high negative association between the fitness value and the plasticity of leaf length was obtained, indicating a cost of plasticity. The selection gradient for leaf angle was weak and significant only in the woodland understory. Genetic correlations between trait expressions in contrasting light environments were negative in sign and low in magnitude, implying a significant genetic variation for plasticity in these leaf traits. Furthermore, leaf length and leaf width were found to be genetically positively coupled, which indicates that there is a potential for these two traits to evolve toward their optimal phenotypic values even faster than would be expected if they were genetically independent.     

亚热带常绿阔叶林不同土壤和林冠环境下蝴蝶花的克隆可塑性

对黄壤和喀斯特土壤两种土壤类型的森林内部和林窗中蝴蝶花的生长指标进行调查,研究异质生境下蝴蝶花的克隆可塑性.结果表明:蝴蝶花在黄壤生境中的分株高度、分株基径和根茎直径均高于喀斯特土壤生境,而分株密度低于喀斯特土壤生境.光照显著影响分株密度,在同一光照条件下土壤质地对分株密度没有显著影响.喀斯特森林内部蝴蝶花的花蕾数及开花数为0,这可能解释为光照和土壤质地双重压力下蝴蝶花对资源摄取和能量分配的权衡与生长维持.在一定程度上,蝴蝶花在黄壤生境中通过增强单个分株的竞争力、减少个体数量来适应环境,趋向K对策,在喀斯特生境中则通过增加个体数量、维持分株竞争力来适应环境,趋向r对策.     

The evolution of phenotypic plasticity in spatially structured environments: implications of intraspecific competition, plasticity costs and environmental characteristics

We model the evolution of reaction norms focusing on three aspects: frequency-dependent selection arising from resource competition, maintenance and production costs of phenotypic plasticity, and three characteristics of environmental heterogeneity (frequency of environments, their intrinsic carrying capacity and the sensitivity to phenotypic maladaptation in these environments). We show that (i) reaction norms evolve so as to trade adaptation for acquiring resources against cost avoidance; (ii) maintenance costs cause reaction norms to better adapt to frequent rather than to infrequent environments, whereas production costs do not; and (iii) evolved reaction norms confer better adaptation to environments with low rather than with high intrinsic carrying capacity. The two previous findings contradict earlier theoretical results and originate from two previously unexplored features that are included in our model. First, production costs of phenotypic plasticity are only incurred when a given phenotype is actually produced. Therefore, they are proportional to the frequency of environments, and these frequencies thus affect the selection pressure to avoid costs just as much as the selection pressure to improve adaptation. This prevents the frequency of environments from affecting the evolving reaction norm. Secondly, our model describes the evolution of plasticity for a phenotype determining an individual's capability to acquire resources, and thus its realized carrying capacity. When individuals are distributed randomly across environments, they cannot avoid experiencing environments with intrinsically low carrying capacity. As selection pressures arising from the need to improve adaptation are stronger under such extreme conditions than under mild ones, better adaptation to environments with low rather than with high intrinsic carrying capacity results.     

Developmental plasticity in plants: implications of non-cognitive behavior

There has been a surge of interest in phenotypic plasticity in the last two decades. Most studies, however, are being carried out within relatively narrow disciplinary frameworks. Consequently, researchers differ not only in their scientific agenda; they often use different terminologies and conceptual frameworks even when studying the very same phenomena. The diversity of approaches has often generated parallel bodies of theory on subjects that can be best understood in broader interdisciplinary terms. This special issue points out the differences between the concepts and questions that are characteristic of various approaches. Bridging all gulfs may be impossible and not necessarily desirable, yet, awareness of the varied approaches should be instrumental in promoting interdisciplinary advances. It is the contribution to such awareness that is the major purpose of this special issue, and for this reason it deals with molecular, physiological, ecological and evolutionary approaches to the study of developmental plasticity. So as to focus the discussion, six topics have been selected, ranging from the fundamental essence of developmental plasticity to its implications to ecology and evolution. These topics were considered by scholars who were chosen for the diversity of their research, not only their expertise. Rather than a comprehensive body of theory, the current issue thus seeks the diversity of opinions on the discussed topics. It is hoped that the confrontation, in its original Latin sense, which includes bringing together and discussion, of scholars who are studying these phenomena at very different levels and from different points of view will generate new insights and promote future interdisciplinary research.     

Phenotypic plasticity of abdominal pigmentation in Drosophila kikkawai: multiple interactions between a major gene, sex, abdomen segment and growth temperature

Drosophila kikkawai is known to be polymorphic for a single autosomal locus controlling abdomen pigmentation in females. Two strains homozygous at this locus (Abdomen pigmentation, Abp) were established from a polymorphic Indian population: one was homozygous (DD) for the dark allele, the other (LL) for the light allele. A Mendelian analysis of crosses at 25°C confirmed the occurrence of a major locus, with dominance of the D allele. Phenotypic variation of pigmentation according to growth temperature was then analyzed in DD and LL male and female flies, and in reciprocal F1. A slight difference was found between reciprocal F1 females from a dark mother were darker but not at all temperatures. In females, the D allele exhibited an antero‐posterior gradient of increasing expression from segment 27, with dominance over L and an increased expression at low temperatures. In males, abdomen pigmentation was uniformly light in segments 25, the D allele being repressed by the sex genotype. In segment 6, the D allele was expressed but only at low temperatures, and was either recessive to L or codominant. Phenotypic plasticity that is, amount of change induced by different growth temperatures, was variable according to genotype and segment. It always corresponded to a darkening of the fly at lower temperatures, but was generally much less than in D. melanogaster. In D. kikkawai, climatic adaptation might occur more by changing the frequency of the D allele than by phenotypic plasticity. This revised version was published online in July 2006 with corrections to the Cover Date.     

Patterns of genotypic variation and phenotypic plasticity of light response in two tropical Piper (Piperaceae) species

Patterns of phenotypic plasticity and genotypic variation in light response of growth and photosynthesis were examined in two species of rain forest shrub that differ in ecological distribution within the forest. We further examined correlations among photosynthetic and growth traits. We hypothesized that the pioneer species, Piper sancti-felicis, would display greater phenotypic plasticity than the shade-tolerant species, Piper arieianum. We further proposed that, in both species, genotypic effects would be more apparent in growth-related traits than photosynthetic traits due to more concentrated selection pressure on gas-exchange traits. P. sancti-felicis did not demonstrate greater phenotypic plasticity of light response. Although many of the traits measured had significant genotype effects, neither species showed any significant effects of genotype on light response of photosynthesis, suggesting little genetic variation for this trait within populations. A principal components analysis clearly illustrated both species and light effects, with the treatments dividing neatly along the axis of the first principal component and the species separating along the second principal component axis. Results indicated general similarities between the species in their trait correlation structure and level of integration among traits, but characteristic differences were observed in the patterns of change between low and high light. Both species had more correlations than expected within groups of growth-related or photosynthetic traits; strong correlations of traits between these two groups were underrepresented. The similar pattern of genetic variation and phenotypic integration observed in these two congeners may be due more to their close phylogenetic relation than to their ecological distributions.     

Pupal period and adult size in Drosophila melanogaster: a cautionary tale of contrasting correlations between two sexually dimorphic traits

Sexual dimorphism (SD) is widespread, reflecting a resolution of genetic conflicts arising from sex-specific differences in selection. However, genetic correlations among traits may constrain the evolution of SD. Drosophila melanogaster exhibits SD for pupal period (males longer) and adult weight (females heavier). This negative inter-sex covariance between the traits contrasts with a significant intra-sex positive genetic correlation (r(g) = 0.95) estimated using lines selected for fast larval development. Path analysis indicated that within sexes the selection regime indirectly reduced adult weight which in turn reduced pupal period. A hypothesis is proposed for the evolution of SD whereby the trait 'pupal period' is divided into 'intrinsic' (correlated with body size) and 'ecological' (uncorrelated with body size) components, and (the larger) females eclose earlier than males size via a shortening of the ecological component, thus achieving the advantage of provisioning eggs prior to sexual maturity. This hypothesis avoids invoking successful 'incompatible antagonistic selection'.     

Flowering time plasticity in Arabidopsis thaliana: a reanalysis of Westerman & Lawrence (1970)

Environmental variation in temperature can have dramatic effects on plant morphology, phenology, and fitness, and for this reason it is important to understand the evolutionary dynamics of phenotypic plasticity in response to temperature. We investigated constraints on the evolution of phenotypic plasticity in response to a temperature gradient in the model plant Arabidopsis thaliana by applying modern analytical tools to the classic data of Westerman & Lawrence (1970). We found significant evidence for two types of constraints. First, we detected numerous significant genetic correlations between plastic responses to temperature and the mean value of a trait across all environments, which differed qualitatively in pattern between the set of ecotypes and the set of mutant lines in the original sample. Secondly, we detected significant costs of flowering time plasticity in two of the three experimental environments, and a net pattern of selection against flowering time plasticity in the experiment overall. Thus, when explored with contemporary methods, the prescient work of Westerman & Lawrence (1970) provides new insights about evolutionary constraints on the evolution of plasticity.     

Effects of four generations of density-dependent selection on life history traits and their plasticity in a clonally propagated plant

Life history evolution of many clonal plants takes place with long periods of exclusively clonal reproduction and under largely varying ramet densities resulting from clonal reproduction. We asked whether life history traits of the clonal herb Ranunculus reptans respond to density-dependent selection, and whether plasticity in these traits is adaptive. After four generations of exclusively clonal propagation of 16 low and 16 high ramet-density lines, we studied life history traits and their plasticities at two test ramet-densities. Plastic responses to higher test-density consisted of a shift from sexual to vegetative reproduction, and reduced flower production, plant size, branching frequency, and lengths of leaves and internodes. Plants of high-density lines tended to have longer leaves, and under high test-density branched less frequently than those of low-density lines. Directions of these selection responses indicate that the observed plastic branching response is adaptive, whereas the plastic leaf length response is not. The reverse branching frequency pattern at low test-density, where plants of high-density lines branched more frequently than those of low-density lines, indicates evolution of plasticity in branching. Moreover, when grown under less stressful low test-density, plants of high-density lines tended to grow larger than the ones of low-density lines. We conclude that ramet density affects clonal life-history evolution and that under exclusively clonal propagation clonal life-history traits and their plasticities evolve differently at different ramet densities.     

Size and architectural traits as ontogenetic determinants of fitness in a phenotypically plastic annual weed (Amaranthus albus)

Abstract Plasticity of size and architectural traits, and their importance to reproductive fitness, were estimated in relation to nutrient availability in the annual Amaranthus albus . Seeds from seven field-collected genotypes were used to rear a first generation under uniform conditions. Seed families (inbred lines) from four first-generation plants per genotype were used to rear a second generation in a glasshouse for 10 weeks. Nine plants per family were regularly fertilized, while nine others were unfertilized. Size was estimated at 5, 8, and 10 weeks; architectural traits were recorded at 8 weeks (no. branches) and 10 weeks (no. branches and branch length). Fitness was assessed by the number of seeds per plant. Because traits were intercorrelated, size-scaled architectural traits were generated and allometric analyses performed. Genetic variation was analyzed for the second generation among inbred lines of the original genotypes, and among families within each genotype. For fertilized and unfertilized groups, early size (volume occupied at 5 weeks) and branch length per mass were significant determinants of fitness. The number of branches per size explained a smaller proportion of the variance in fitness. Genotype by treatment interaction was apparent for some traits, indicating genetic variation for plasticity, but plasticity of size did not change over ontogeny. Significant effects of genotype on size and architectural traits, and fitness, were detected mostly in the unfertilized group. Thus, selection is most likely to differentiate among genotypes in nutrient-poor environments. Lengthening of multiple branches increases seed output throughout the season, and genotypes with longer branches per unit mass have a selective advantage. Because early size is correlated with seed output, ontogenetic plasticity in response to improved soil resources allows opportunistic increases in fitness.     

The impact of plasticity and maternal effect on the evolution of leaf resin production in Diplacus aurantiacus

Our previous quantitative genetic study of leaf resin production in Diplacus aurantiacus revealed large environmental and maternal effects on variation in resin production, which suggests the possibility of a genotype×environment interaction for this trait when plants grow in heterogeneous environments. Our objectives in this study were to observe the genetic variation in plasticity of resin production under field and chamber conditions, compare phenotypic correlations of resin content with growth traits under these two environmental conditions, and distinguish the possible basis of the maternal effect on resin production using parents and half-sib progeny. A significant genotype×environment interaction (P<0.0001) in leaf resin production was found, which suggests a potential for the evolution of plasticity of these secondary metabolites under heterogeneous environments. The phenotypic correlation between resin content and growth rate also exhibited plasticity. In addition, the resin content of dam half-sib families grown in the chamber had a closer relationship with their maternal parents in the field (r=0.65, P=0.059) than in the chamber (r=0.39, P=0.34), suggesting an environmentally based maternal effect on the secondary chemicals. We suggest that the maternal environmental effect may act as a contributor to plasticity of resin production and, while it may not diminish the appearance of the genotype×environment interaction, the heritable variation of plasticity of resin production may be confounded.     

The role of plant traits and their plasticity in the response of pasture grasses to nutrients and cutting frequency

MethodsMonocultures of the grass species were grown in a fully factorial block design combining plant species, cutting frequency and N supply as factors.ConclusionsVariations in ANPP of species in response to an increase in cutting frequency and a decrease in N supply are controlled by a group of traits, rather than by one individual trait. Incorporating plasticity of the individual traits into these trait combinations was the key to explaining species'' productivity responses, accounting for up to 89 % of the total variability in response to the changes in N supply.     

Phenotypic plasticity of body size in Drosophila: effects of a daily periodicity of growth temperature in two sibling species

Abstract. Variation of wing and thorax length under thermoperiodic growth conditions was analysed in four strains of two sibling species, Drosophila melanogaster and D. simulans , from two European localities. Results were compared to those obtained with constant temperatures ranging from 12 to 31 °C.
Under constant temperatures the data basically confirmed previous results: concave reaction norms for wing and thorax length; a monotonically decreasing norm for wing : thorax ratio; and an increasing norm for sex dimorphism (female : male ratio). Phenotypic variability was maximum at extreme temperatures and minimum at middle ones. Slight differences were observed according to the geographical origin: the difference between strains from Bordeaux (France) and Cordoba (Spain) was maximum at low temperatures but disappeared at about 28 °C.
According to the temperatures chosen, alternating thermal regimens had either no effect or produced a significant size reduction, probably reflecting a periodic stress. The magnitude of this effect was proportional to the amplitude of the thermoperiod but not to the quality (cold or heat) of the stress. In a similar way, the wing : thorax ratio was either not modified or reduced significantly, indicating that wing length was relatively more affected than thorax length by alternating thermal regimens. Sex dimorphism also showed either no change or a significant increase, indicating that males were relatively more reactive than females to alternating conditions. Finally, regimens of broad amplitudes increased the phenotypic variability, again an indication of stressful effects. All these observations should be taken into account when analysing phenotypic variability in nature and trying to understand natural selection in wild-living populations.     

江南油杉种实性状变异及其与环境因子的关系

该研究以广西低海拔盆地向云贵高原过渡的连续地理区域内江南油杉(Keteleeria fortunei var.cyclolepis)林为对象,采用方差分析、多重比较、相关分析、逐步回归分析等方法,对4省(区)9个江南油杉天然种群的16个种实表型性状进行比较研究,探讨种群间种实表型性状多样性及其与地理环境因子的关系。结果表明:江南油杉种实表型性状在种群间存在丰富变异,16个种实性状在种群间均达到显著或极显著差异;9个种群中望谟种群变异最大(2.54%),16个种实表型性状中种子宽变异最大(9.04%),种群间种子性状的变异高于球果的变异,说明球果性状变异稳定性高;种实部分表型性状间呈显著或极显著相关,其中种子宽、种子厚、种子长宽比、种鳞长宽比、苞鳞长宽比是较为关键的性状,能在一定程度上反映种实质量;种子性状受地理环境因子的影响比球果大,表现为西部种源种子更趋于圆形,饱满,种翅较长,较宽;地理环境因子中,海拔与多数种实性状间呈显著相关性,是种实表型性状变异的主要来源,因此,海拔是影响种实表型性状变异的主要因素。该研究结果对进一步保护、利用江南油杉种质资源具有一定的指导意义。     

Effects of nitrogen supply on photosynthetic and anatomical changes in current-year needles of <Emphasis Type="Italic">Pinus koraiensis</Emphasis> seedlings grown under two irradiances

We investigated the responses of photon-saturated photosynthesis rate (P _sat) and its simultaneous acclimation of anatomy and nitrogen use patterns of current needles of Korean pine (Pinus koraiensis) seedlings grown under factorial combinations of two nitrogen levels and irradiances. Although N supply resulted in a significant increase of N content in needles under both irradiances, the increase of P _sat tended to be suppressed only in shade (S). The significant increase of P _sat in full sunlight (O) was associated with the increase of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and chlorophyll (Chl) contents. In contrast, small increase of Chl content and no increase of RuBPCO content were found in S (90 % cut of full irradiance), which would result in a small increase of P _sat. This result suggests that extra N is stocked in needles under shade for the growth in next season. With N supply, a significant decrease of specific leaf area (SLA) was detected only in O. This decrease of SLA was due to the increase of density of needle. Furthermore, the increase of needle density was not due to the increased number and size of mesophyll cells, but the increased density of each mesophyll cell. Therefore, although SLA changed in O, the change did not involve anatomical adaptation to use increased N effectively, at least observable by light microscopy. Hence, even though the SLA would change, N deposition will improve the photosynthetic capacity of Korean pine seedlings, not through the development of needle anatomy but through improvement of the allocation of N in both irradiances.     

Inter- and intraspecific variation in the germination response to light quality and scarification in grasses growing in two-phase mosaics of the Chihuahuan Desert

BACKGROUND AND AIMS: In many locations, plants are faced with adjacent, contrasting environments, and the between-species differential evolution of life history traits can be interpreted as an evolutionary response to this environmental heterogeneity. However, there has been little research on the intraspecific variability in these attributes as a possible evolutionary response of plants. METHODS: In the two-phase mosaic of the Chihuahuan Desert (adjacent patches with contrasting resource availability), analyses were carried out of the germination response to the scarification and light quality to which grass seeds growing on these patches are exposed (open and closed habitats). KEY RESULTS: Species that grow in open habitats exhibited a higher germination success than those from closed habitats after scarification. At both the inter- and intraspecific level, there were differences in the germination percentage and in the germination speed in response to light quality. Intraspecific variation in the species from the closed habitat (Pleuraphis mutica and Trichloris crinita) and in Chloris virgata (which grows in both habitats) was due to genetic variation (the family factor was significant), but there was no genetic variation in phenotypic plasticity (non-significant interaction between family and light quality). In contrast, for the species that grows only in the open habitat (Dasyochloa pulchella), the family did not have a significant effect, but there was genetic variation in the phenotypic plasticity (significant interaction between family and light quality). CONCLUSIONS: In C. virgata, P. mutica and T. crinita, natural selection could be favouring those genotypes that responded better in each light environment, but it is not possible that the natural selection resulted in different optimal phenotypes in each habitat. On the contrary, in D. pulchella, selection could have reduced the genetic variation, but there is the possibility of the evolution of reaction norms, resulting in the selection of alternative phenotypes for each habitat.     

福建茶树害螨发生动态及其与茶树品种理化特性的相关性分析

用5点采叶取样法调查了茶树害螨发生特点。结果表明:各茶树品种的害螨盛发高峰期一般都出现在8月底到9月初;对2005年5品种茶树上害螨螨情指数高峰期进行方差分析,结果表明均达到极显著差异(F=4.141,P=0.0054)。对不同茶树品种的叶片理化指标与螨情指数进行相关分析,结果表明:叶片厚度与相应品种的螨情指数之间呈一定的负相关关系(F=0.3903,r=-0.4040);不同茶树品种的叶片下表皮及其角质层厚度和相应品种的螨情指数之间呈一定的正相关关系(F=2.40,r=0.7389);不同茶树品种叶片的咖啡因和GC含量与害螨螨情指数呈负相关,相关关系接近显著水平(F=3.44,r=-0.7309和F=5.96,r=-0.8156)。