Niche construction through phenological plasticity: life history dynamics and ecological consequences

The ability of an organism to alter the environment that it experiences has been termed 'niche construction'. Plants have several ways whereby they can determine the environment to which they are exposed at different life stages. This paper discusses three of these: plasticity in dispersal, flowering timing and germination timing. It reviews pathways through which niche construction alters evolutionary and ecological trajectories by altering the selective environment to which organisms are exposed, the phenotypic expression of plastic characters, and the expression of genetic variation. It provides examples whereby niche construction creates positive or negative feedbacks between phenotypes and environments, which in turn cause novel evolutionary constraints and novel life-history expression.     

Phenotypic homogeneity of two intertidal snails across a wave exposure gradient in South Australia

Dislodgement by the large drag forces imparted by breaking waves is an important cause of mortality for intertidal snails. The risk of drag-induced dislodgement can be reduced with: (1) a smaller shell of lower maximum projected surface area (MPSA); (2) a streamlined shell shape characterized by a squatter shell; and/or (3) greater adhesive strength attained through a larger foot area or increased foot tenacity. Snails on exposed coasts tend to express traits that increase dislodgement resistance. Such habitat-specific differences could result from direct selection against poorly adapted phenotypes on exposed shores but may reflect gastropod adaptation to high wave action achieved through phenotypic plasticity or genetic polymorphism. With this in mind, we examined the size, shape and adhesive strength of populations of two gastropod species, Austrocochlea constricta (Lamarck) and Nerita atramentosa (Reeve), from two adjacent shores representing extremes in wave exposure. Over a 5 day period, maximum wave forces were more than 10 times greater on the exposed than sheltered shore. Size-frequency distributions indicate that a predator consuming snails within the 1.3-1.8 cm length range regulates sheltered shore populations of both snail species. Although morphological scaling considerations suggest that drag forces should not place physical limits on the size of these gastropods, exposed shore populations of both snails were small relative to the maximum size documented for these species. Therefore, selective forces at the exposed site might favour smaller individuals with increased access to microhabitat refuges. Unexpectedly, however, neither snail species exhibited between-shore differences in shape, foot area or foot tenacity, which are likely to have adaptive explanations. Hence, it is possible that these snails are incapable of adaptive developmental responses to high wave action. Instead, the homogeneous and wave-exposed nature of Australia's southern coastline may have favoured the evolution of generalist strategies in these species.     

Allozyme and shell variation in two marine snails (Littorina, Prosobranchia) with different dispersal abilities

As adults Littorina littorea and L. saxatilis have overlapping distributions, both on the shore and on a geographical scale, this implies that they are affected by similar selective forces to a large extent. The two species have, however, different reproductive strategies. Littorina littorea has a planktonic larval stage lasting 4–6 weeks, whereas L. saxatilis has direct development. This influences dispersal rates of the two species, and gene flow and effective population size are both assumed to be much smaller in L. saxatilis than in L. littorea. Intraspeciflc variation in morphological and allozyme characters differs between the two species. Littorina littorea is homogeneous for both types of characters, both within and between populations, while populations of L. saxatilis may show pronounced differences in shell characters with micro-environment changes, and allozyme variation is markedly higher in this species too. The polymorphism in shell characters of L. saxatilis and the monomorphism of L. littorea, may be expected from theory as a consequence of the different dispersal strategies. Also the lack of divergence between geographically separate populations of L. littorea, but the high degree of between population differentiation in L. saxatilis, could be attributed to the high and low dispersal rates, respectively. However, the larger allozyme variation within subpopulations of L. saxatilis compared to that of L. littorea, is surprising as the former species has a smaller effective population size than the latter. Furthermore, there is no correlation between dispersal ability and overall allozyme heterozygosity, when 10 additional species of Littorina are included in the comparison. It is suggested that different evolutionary backgrounds of different species will possibly explain this, as neither selectionist nor neutralist models alone account for the observation. That is, an earlier passage through a population bottleneck may, for example, influence the genetic variation for a long period of time.     

Autotomy-induced life history plasticity in band-legged ground cricket Dianemobius nigrofasciatus

Crickets can autotomize their limbs when attacked by predators. This enables them to escape death, but imposes a short-term cost on their escape speed and a long-term cost on their future mating ability. Therefore, adaptive response compensated for the cost of autotomy might be advantageous for autotomized individuals. In the present study, we examined whether autotomy induced life history plasticities compensating for the future cost in the band-legged ground cricket Dianemobius nigrofasciatus . Life history traits of D. nigrofasciatus were compared between autotomized and intact individuals. The developmental time and head width of the individuals that were autotomized as fourth instar nymphs were significantly shorter and smaller, respectively, than those of intact individuals. However, the adult longevity, number of eggs laid and oviposition schedule did not vary between autotomized and intact individuals. In addition, there was no difference between individuals autotomized at the fourth instar and adult stages in these three traits. Early maturation in the autotomized individuals might be advantageous through reducing the risk of predation owing to the shorter period in nymphal stages. The cost of small body size in the autotomized females might not be so great because of no significant difference in fecundity between autotomized and intact individuals. However, the cost of small body size was unclear in the autotomized males because in general larger males were preferred by females. These results indicated autotomy-induced life history that might reduce the cost of autotomy.     

Geographic variation and plasticity to water and nutrients in Pelargonium australe

Here, patterns of phenotypic plasticity and trait integration of leaf characteristics in six geographically discrete populations of the perennial herb Pelargonium australe were compared. It was hypothesized that populations would show local adaptation in trait means, but similar patterns of plasticity and trait integration. Further, it was questioned whether phenotypic plasticity was positively correlated with environmental heterogeneity and whether plasticity for water-use traits in particular was adaptive. Seedlings were grown in a glasshouse at six combinations of water and nutrient availability. Leaf anatomical, morphological and gas exchange traits were measured. High amounts of plasticity in leaf traits were found in response to changes in growth conditions and there was evidence of local adaptation among the populations. While there were significant correlations between plasticity and environmental heterogeneity, not all were positive. Notably, patterns of plasticity and trait integration varied significantly among populations. Despite that variation, some of the observed plasticity was adaptive: fitness was correlated with conservative water use when water was limiting. Pelargonium arrived in Australia approximately 5 million yr ago. It is concluded here that high amounts of plasticity, in some cases adaptive, and weak integration among traits may be key to the spread and success of this species.     

Phenotypic variation in colour pattern and seasonal plasticity in Eristalis hoverflies (Diptera: Syrphidae)

Abstract.

• 1 An examination of phenotypic variation in colour pattern was carried out on four Eristalis hoverfly species using museum material.
• 2 The amount of phenotypic variation varied substantially among the species with E.arbustorum being the most variable. The other species showed a wide colour pattern range but less variation within that range (E.abusivus and E.nemorum), or a narrow range of colour variation (E.horticola).
• 3 Sexual colour dimorphism was apparent in all four species, but most pronounced in E.abusivus and E.nemorum.
• 4 There were good phenotype-season relationships shown by both sexes in all species, except for female E.abusivus and E.nemorum, with paler insects being more abundant during the warmer summer months.
• 5 Female, but not male, E.arbustorum collected at inland sites were on average paler than those collected at coastal sites. This observation is considered with respect to temperature during the developmental stages.
• 6 The function of colour plasticity in hoverflies is discussed with reference to the need to maintain optimal thermal conditions for activity.

     

Genetic and maternal influences on life history plasticity in milkweed bugs (Oncopeltus): response to temperature

This study was designed to examine life history flexibility arising from phenotypic plasticity in response to temperature and from maternal effects in response to reproductive diapause in a temperate zone population of the milkweek bug (Oncopeltus fasciatus). We employed a split-family, first-cousin, full-sib design with siblings reared at different temperatures in order to quantify phenotypic plasticity, maternal effects, and variation for each. The following traits were analyzed: development time, age at first reproduction, longevity, early-life fecundity, and wing length. We found both life history plasticity and maternal effects on life history traits which tend to enhance the colonizing ability of offspring born to mothers that have undergone reproductive diapause. We were unable to demonstrate additive genetic variation for plasticity for any of the traits, while for development time and wing length we found variation due to non-additive genetic or common-environmental sources. We were also unable to demonstrate additive genetic variation for maternal effects, although variation may exist at low levels that are difficult to detect using cousin-families. The apparent lack of variation in this population would constrain evolution of life history flexibility even though considerable flexibility exists in the phenotype.     

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.     

Rapid parallel evolution of standing variation in a single,complex, genomic region is associated with life history in steelhead/rainbow trout

Rapid adaptation to novel environments may drive changes in genomic regions through natural selection. Such changes may be population-specific or, alternatively, may involve parallel evolution of the same genomic region in multiple populations, if that region contains genes or co-adapted gene complexes affecting the selected trait(s). Both quantitative and population genetic approaches have identified associations between specific genomic regions and the anadromous (steelhead) and resident (rainbow trout) life-history strategies of Oncorhynchus mykiss. Here, we use genotype data from 95 single nucleotide polymorphisms and show that the distribution of variation in a large region of one chromosome, Omy5, is strongly associated with life-history differentiation in multiple above-barrier populations of rainbow trout and their anadromous steelhead ancestors. The associated loci are in strong linkage disequilibrium, suggesting the presence of a chromosomal inversion or other rearrangement limiting recombination. These results provide the first evidence of a common genomic basis for life-history variation in O. mykiss in a geographically diverse set of populations and extend our knowledge of the heritable basis of rapid adaptation of complex traits in novel habitats.     

Phenotypic plasticity to light of two congeneric trees from contrasting habitats: Brazilian Atlantic Forest versus cerrado (savanna)

The Brazilian Atlantic Forest is a typically multi-layer tropical forest, while cerrado (savanna) is a patchy habitat with different physiognomy. Despite these differences, both habitats have high light heterogeneity. Functional traits of Dalbergia nigra and D. miscolobium from the Atlantic Forest and cerrado, respectively, were evaluated under shade (25% of full sunlight) and full sunlight in a nursery experiment. We hypothesised that both species should benefit from high phenotypic plasticity in relation to light. Plasticity was estimated using the relative distance phenotypic index (RDPI). D. miscolobium had lower shoot growth under both light conditions, suggesting it has low competitive capacity in the forest environment, which could explain its limited ability to expand over areas of Atlantic Forest. The studied species exhibited photoprotection strategies under high light and improved light capture under low light. Stomatal conductance, ETR(max) (maximum electron transport rate), PPFD(sat) (saturating photosynthetically active photon flux density), chlorophyll and carotenoid content had higher RDPI than stem morphological traits. Although both species showed considerable phenotypic plasticity, D. miscolobium had higher RDPI for eight of 11 evaluated traits. This high plasticity could be one of the factors that explain the occurrence of this species in a wide range of environmental conditions, from open grassland to dense woodlands, and it could also reflect its adaptation to high light. D. nigra also had considerable plasticity and good growth performance in both shade and full sunlight, but its absence in areas of cerrado suggests that factors other than light limit its occurrence in these habitats.     

Interclonal differences, plasticity and trade-offs of life history traits of Cyperus esculentus in relation to water availability

Cyperus esculentus is an exotic clonal (or pseudoannual) weed in Japan, and its range is steadily increasing. To investigate its interclonal variation and phenotypic plasticity in response to water availability, five clones of C. esculentus , collected from different sites in Japan, were grown singly in pots placed outdoors under dry and wet conditions. All the traits examined showed considerable variation among the five clones. However, two clones from Tochigi were similar to each other; thus, they might have originated from the same founder population. The clone from Ibaraki was quite different from the others. Therefore, it is suggested that the Japanese populations of C. esculentus might have resulted from multiple introductions of genotypes from geographically separated and, hence, genetically differentiated, source populations. All the clones also showed considerable plasticity in response to water availability. Clones with a larger ramet number had a greater plasticity, whereas tuber size was invariant across water treatments. Highly plastic traits had generally low interclonal variation in plasticity. All the clones had high productivity and produced more ramets and tubers under wet conditions than under dry conditions. Moreover, water availability could partially regulate the mode of its reproduction; wet conditions favored tuber production (vegetative propagation) while dry conditions favored sexual reproduction. A number of trade-offs occurred between the traits of clonal growth, storage and sexual reproduction, indicating that allocation among the competing functions/organs is mutually exclusive in plants. The results obtained here suggest that C. esculentus is more likely to invade wet habitats than dry habitats.     

Phenotypic plasticity and the evolution of trade-offs: the quantitative genetics of resource allocation in the wing dimorphic cricket, Gryllus firmus

In the wing dimorphic sand cricket, Gryllus firmus, there is a pronounced trade-off between flight capability and fecundity. This trade-off is found both between morphs and within the macropterous morph, in which fecundity is negatively correlated with the mass of the principle flight muscles, the dorso-longitudinal muscles (DLM). In this paper, we examine how this trade-off is affected by a reduction in food and its genetic basis. We find that the relative fitness of the two wing morphs is not changed although both fecundity and DLM mass are decreased. A quantitative genetic analysis shows that the trade-off function is genetically variable but that most of the variation occurs in the intercept rather than the slope of the function. Analysis further indicates a very high genetic correlation between environments (food ration) supporting the hypothesis of a strong functional constraint between reproduction and flight capability.     

Light quantity and quality induce shade-avoiding plasticity in a marine macroalga

Light-induced plasticity in plant morphology is considered adaptive in terrestrial habitats that vary in light, but remains unexplored for marine habitats. This is despite similar modes of growth, development and photosynthetic equipment in terrestrial and marine photoautotrophs and similarly dynamic light environments. We tested whether manipulations of light quantity and quality induce morphological plasticity in the marine macroalga, Asparagopsis armata. Using multivariate analyses (principal components analyses and multivariate analyses of covariance), we show that correlated morphological traits underlie a fundamental growth strategy characterized by the production of phalanx and guerrilla phenotypes in environments that mimic light and shade respectively. This foraging response is not under simple genetic or environmental control, but influenced by interactions between genotype and environment. Evidence of plasticity and genetic variation in plasticity in a marine modular organism generates additional, testable hypotheses on the ecological consequences of variation in growth form that may further explain the evolution of plasticity.     

Seasonal plasticity in life history traits: growth and development in Polygonia c-album (Lepidoptera: Nymphalidae)

The potentially multivoltine comma butterfly, Polygonia c-album L., hibernates in the adult stage. The adult seasonal morph is demonstrated to be a good indicator of whether an individual has entered reproductive diapause or is developing directly to sexual maturation. This fact, and the assumption that a short development time is not equally important to all categories of individuals, was used to test predictions on variation in life-history traits among categories (morphs and sexes) and environments (temperature and photoperiod) at the level of individuals and to some extent families and populations (the univoltine Stockholm population and the partially bivoltine Oxford population). Individuals developing to adults in a short time were expected to be smaller and lighter as a result of a basic trade-off between the two traits. Development times varied in accordance with predictions, but in most cases this was due to plastic growth and development in both the larval and pupal stages rather than through variation in size or weight, i.e. size was a highly canalized trait. This suggests a relationship between plasticity and canalization and a strong potential for plasticity to shield life-history traits from selection. Individuals regulated development times also within developmental pathways, in response to photoperiods indicating the progression of the season. These and other results suggest that development times are not normally minimized in temperate butterflies unless this is enforced by direct development and protandry. There is thus scope for a high degree of adaptive plasticity in growth- and developmental rates which may devalue the basic trade-offs assumed by life-history theory and account for inconsistencies with its predictions.     

Phenotypic plasticity and specialization in clonal versus non-clonal plants: A data synthesis

Reproductive strategies can be associated with ecological specialization and generalization. Clonal plants produce lineages adapted to the maternal habitat that can lead to specialization. However, clonal plants frequently display high phenotypic plasticity (e.g. clonal foraging for resources), factors linked to ecological generalization. Alternately, sexual reproduction can be associated with generalization via increasing genetic variation or specialization through rapid adaptive evolution. Moreover, specializing to high or low quality habitats can determine how phenotypic plasticity is expressed in plants. The specialization hypothesis predicts that specialization to good environments results in high performance trait plasticity and specialization to bad environments results in low performance trait plasticity. The interplay between reproductive strategies, phenotypic plasticity, and ecological specialization is important for understanding how plants adapt to variable environments. However, we currently have a poor understanding of these relationships. In this study, we addressed following questions: 1) Is there a relationship between phenotypic plasticity, specialization, and reproductive strategies in plants? 2) Do good habitat specialists express greater performance trait plasticity than bad habitat specialists? We searched the literature for studies examining plasticity for performance traits and functional traits in clonal and non-clonal plant species from different habitat types. We found that non-clonal (obligate sexual) plants expressed greater performance trait plasticity and functional trait plasticity than clonal plants. That is, non-clonal plants exhibited a specialist strategy where they perform well only in a limited range of habitats. Clonal plants expressed less performance loss across habitats and a more generalist strategy. In addition, specialization to good habitats did not result in greater performance trait plasticity. This result was contrary to the predictions of the specialization hypothesis. Overall, reproductive strategies are associated with ecological specialization or generalization through phenotypic plasticity. While specialization is common in plant populations, the evolution of specialization does not control the nature of phenotypic plasticity as predicted under the specialization hypothesis.