FUNCTIONAL MORPHOLOGY AND FEEDING OF DONAX SERRA ROODING AND DONAX SORDIDUS HANLEY (BIVALVIA: DONACIDAE)

The functional morphology of Donax serra and D. sordidus fromSouth African beaches was examined, and comparative measurementsmade of the rate of water filtration. The two species differin the structure and ciliation of the ctenidia, the number andspacing of the ridges on the labial palps, the direction ofciliary currents on the surface of the mantle and visceral mass,and the length and coiling of the mid-gut. The maximum ratesof water filtration recorded showed similar relationships tosize for the two species, but in D. sordidus the rate declinedrapidly with time during the observations. These differencesindicate differences in feeding strategy related to differencesin distribution and behaviour of the two species on beaches.     

PRELIMINARY ANALYSIS OF QUANTITATIVE GENETICS AND PHENOTYPIC PLASTICITY IN AULACOSEIRA SUBARCTICA (BACILLAR-IOPHYTA)

Several clones of Aulacoseira subarctica were isolated from Yellowstone, Lewis, and East Rosebud Lakes (Montana, Wyoming). Two to four clones from each lake were grown in batch cultures under three light intensities, 2, 11.4 and 115 μE m⁻² s⁻¹. Clones were conditioned to their light environment for a three-week period. Inoculants from the conditioned clones taken during log phase of growth, were grown until in log phase, then samples were collected. Five randomly chosen valves for 2 replicates of each clone were examined using a scanning electron microscope and captured on film at a magnification of 20,000x. Each image was digitized and quantitative morphometric characters were measured. A preliminary quantitative genetic analysis was performed on selected characters within each light environment. Plasticity of characters within clones across the three light regimes were also examined. The amount of variability found within characters in A. subarctica will be discussed in terms of environmental, genetic, and microenvironmental sources.     

AN INTERPRETATION OF PHENOTYPIC PLASTICITY IN AGROPYRON REPENS (GRAMINAE)

Phenotypic plasticity and genotypic variation were studied in Agropyron repens L. (Beauv.) collected from populations in two grassland communities which differed in the length of time since the last major disturbance. Twenty genotypes were collected from each population. Each genotype was vegetatively propagated, and subjected to six different treatments in a greenhouse. Phenotypic plasticity and genotypic variability were measured as across- and within-treatment standardized variances respectively. Patterns of plasticity were measured by genotype correlations across treatments. The results were presented graphically by the regression method of Garbutt and Zangerl (1983). Analysis of variance revealed significant population, genotype and treatment effects. Significant positive correlations between magnitude and variability of performance were found for all characters. Phenotypic plasticity and magnitude of performance were generally greater in plants collected from the older established field. Evidence for greater specialization in the older population was suggested by negative correlations between performance in the most favorable and least favorable treatments and by greater dissimilarity of genotype response across treatments. A more variable phenotypic response across treatments (i.e., higher plasticity) for plants from the older population may therefore be a consequence of specialization and not an adaptive trait per se.     

LOCAL ADAPTATION AND THE EVOLUTION OF PHENOTYPIC PLASTICITY IN TRINIDADIAN GUPPIES (POECILIA RETICULATA)

Divergent selection pressures across environments can result in phenotypic differentiation that is due to local adaptation, phenotypic plasticity, or both. Trinidadian guppies exhibit local adaptation to the presence or absence of predators, but the degree to which predator‐induced plasticity contributes to population differentiation is less clear. We conducted common garden experiments on guppies obtained from two drainages containing populations adapted to high‐ and low‐predation environments. We reared full‐siblings from all populations in treatments simulating the presumed ancestral (predator cues present) and derived (predator cues absent) conditions and measured water column use, head morphology, and size at maturity. When reared in presence of predator cues, all populations had phenotypes that were typical of a high‐predation ecotype. However, when reared in the absence of predator cues, guppies from high‐ and low‐predation regimes differed in head morphology and size at maturity; the qualitative nature of these differences corresponded to those that characterize adaptive phenotypes in high‐ versus low‐predation environments. Thus, divergence in plasticity is due to phenotypic differences between high‐ and low‐predation populations when reared in the absence of predator cues. These results suggest that plasticity might initially play an important role during colonization of novel environments, and then evolve as a by‐product of adaptation to the derived environment.     

POPULATION STRUCTURE,BIOMASS ALLOCATION,AND PHENOTYPIC PLASTICITY IN MURDANNIA KEISAK (COMMELINACEAE)

Field and glasshouse studies were used to explain differences in plant biomass, shoot length, and reproductive effort in four populations of a wetland annual herb, Murdannia keisak. Populations were chosen from continually thermally disturbed, intermittently thermally disturbed, revegetating, and undisturbed portions of a floodplain forest in South Carolina, USA. Plants in the two thermally disturbed areas were shorter, flowered earlier in the autumn, and produced more and smaller seeds than plants in revegetating and undisturbed sites. Reproductive effort was higher in populations from undisturbed and revegetating sites than in the two thermally disturbed sites. Generally, differences observed in the field were not expressed in the glasshouse plants. Glasshouse experiments suggested that most of the observed among-population differences in size and reproductive effort in the field study were a result of a plastic response to water depth and light. The combination of field and glasshouse data showed that this wetland weed adjusts readily to newly disturbed habitats, thus spreading rapidly and maintaining local dominance.     

GENE FLOW AND SELECTION ON PHENOTYPIC PLASTICITY IN AN ISLAND SYSTEM OF RANA TEMPORARIA

Gene flow is often considered to be one of the main factors that constrains local adaptation in a heterogeneous environment. However, gene flow may also lead to the evolution of phenotypic plasticity. We investigated the effect of gene flow on local adaptation and phenotypic plasticity in development time in island populations of the common frog Rana temporaria which breed in pools that differ in drying regimes. This was done by investigating associations between traits (measured in a common garden experiment) and selective factors (pool drying regimes and gene flow from other populations inhabiting different environments) by regression analyses and by comparing pairwise F_ST values (obtained from microsatellite analyses) with pairwise Q_ST values. We found that the degree of phenotypic plasticity was positively correlated with gene flow from other populations inhabiting different environments (among‐island environmental heterogeneity), as well as with local environmental heterogeneity within each population. Furthermore, local adaptation, manifested in the correlation between development time and the degree of pool drying on the islands, appears to have been caused by divergent selection pressures. The local adaptation in development time and phenotypic plasticity is quite remarkable, because the populations are young (less than 300 generations) and substantial gene flow is present among islands.     

PHENOTYPIC PLASTICITY OF BRANCHING PATTERN IN SUGAR MAPLE (ACER SACCHARUM)

Branching patterns of leader shoots and lower branches were compared in 15 sugar maple trees. The number of first-order branches per second-order branch (R_1:2) differed very significantly between leader shoots and lower branches. Values of R_1:2 averaged 6.47 for leaders and 4.06 for lower branches. Stepwise bifurcation ratios between successive higher orders did not differ significantly. The difference in R_1:2 between leader shoots and lower branches within individual trees is the result of developmental-phenotypic interactions, such that the number of leaf-bearing shoots per support shoot decreases over time. These results indicate that branching patterns within individual trees are not stationary, as has commonly been claimed.     

PHENOTYPIC PLASTICITY AND HETEROCHRONY IN CICHLASOMA MANAGUENSE (PISCES,CICHLIDAE) AND THEIR IMPLICATIONS FOR SPECIATION IN CICHLID FISHES

Cichlid fishes in African rift lakes have undergone rapid speciation, resulting in “species flocks” with more than 300 endemic species in some of the lakes. Most researchers assume that there is little phenotypic variation in cichlid fishes. I report here extensive phenotypic plasticity in a Neotropical cichlid species. I examined the influence of diet on trophic morphology during ontogeny in Cichlasoma managuense. Two groups of full siblings were fed two different diets for eight months after the onset of feeding; thereafter both groups were fed a common diet. Phenotypes that differed significantly at 8.5 months converged almost completely at 16.5 months. If feeding on two different diets is continued after 8.5 months, the phenotypes remain distinct. Differences in diet and possibly in feeding mode are believed to have caused these phenotypic changes. Phenotypic plasticity is described in terms of a qualitative model of heterochrony in which phenotypic change in morphology is explained as retardation of the normal developmental rate. If phenotypic expression of morphology is equally plastic in African cichlid species as it may be in the American cichlids, as exemplified by C. managuense, then taxonomic, ecological, and evolutionary analyses of “species flocks” may be in need of revision. However, Old World cichlids may be less phenotypically plastic than New World cichlids, and this may contribute to the observed differences in speciation rate and degree of endemism.     

PHENOTYPIC PLASTICITY IN DRAPARNALDIA (CHAETOPHORACEAE). II. THE PHYSICAL ENVIRONMENT AND CONCLUSIONS

Eight isolates of Draparnaldia from a variety of freshwater habitats were grown in unialgal, defined culture. The morphological responses of these isolates to different daylengths, light intensities and temperatures are described. The phenotypic expression of Draparnaldia is markedly modified by all three factors. The interaction of these environmental variables was highly significant, and it is not possible, in most cases, to describe or predict the morphological appearance of Draparnaldia by referring to one of these parameters in isolation. The adaptive significance of phenotypic plasticity in Draparnaldia is discussed.     

PHENOTYPIC PLASTICITY IN CHRYSOPERLA: GENETIC VARIATION IN THE SENSORY MECHANISM AND IN CORRELATED REPRODUCTIVE TRAITS

A genetically variable sensory mechanism provides phenotypic plasticity in the seasonal cycle of the Chrysoperla carnea species-complex of green lacewings. The mechanism functions as a switch during the pupal and early imaginal stages to determine aestival reproduction versus aestival dormancy, and it has two major components: (1) response to photoperiod and (2) response to a stimulus(i) associated with the prey of the larvae. Ultimately, the switch is based on the response to photoperiod—an all-or-nothing trait whose variation (long-day reproduction versus a short-day/long-day requirement for reproduction) is determined by alleles at two unlinked autosomal loci. In eastern North America, variation in this component of the switch differentiates two reproductively isolated “species” that are sympatric throughout the region: Chrysoperla carnea, in which both loci are homozygous for the dominant alleles that determine long-day, spring and summer reproduction and thus multivoltinism, and C. downesi, which has a very high incidence of the recessive alleles for the short-day/long-day requirement, and thus univoltine spring breeding. In contrast, geographical populations in western North America harbor variable amounts of within-and among-family genetic variation for the photoperiodic responses and also for the switch's second component—adult responsiveness to the prey of the larvae. The geographic pattern of genetic variation in the two components of the switch indicates that it is a highly integrated adaptation to environmental heterogeneity. Expression of among-family variation in the prey component of the switch is highly dependent on photoperiodic conditions and genotype (it requires a constant long daylength and the recessive short-day/long-day genotype). Thus, we infer that responsiveness to prey evolved as a modifier of the photoperiodic trait. The switch has a significant negative effect on a major determinant of fitness; it lengthens the preoviposition period in nondiapausing reproductives. This negative effect may result in temporal variation in the direction of selection, which helps maintain genetic variability in the switch mechanisms of western populations. Also, the photoperiodic and prey components of the switch are positively correlated with fecundity in nondiapausing reproductives; however, the strong influence of environmental factors—presence or absence of prey—leaves open the question whether the correlated effects on fecundity are expressed in nature.     

GEOGRAPHIC VARIATION AND PHENOTYPIC PLASTICITY OF NUMBER OF TRUNK VERTEBRAE IN SLENDER SALAMANDERS,BATRACHOSEPS (CAUDATA: PLETHODONTIDAE)

To understand the evolutionary significance of geographic variation, one must identify the factors that generate phenotypic differences among populations. I examined the causes of geographic variation in and evolutionary history of number of trunk vertebrae in slender salamanders, Batrachoseps (Caudata: Plethodontidae). Number of trunk vertebrae varies at many taxonomic levels within Batrachoseps. Parallel clines in number occur along an environmental gradient in three lineages in the Coast Ranges of California. These parallel clines may signal either adaptation or a shared phenotypically plastic response to the environmental gradient. By raising eggs from 10 populations representing four species of Batrachoseps, I demonstrated that number of trunk vertebrae can be altered by the developmental temperature; however, the degree of plasticity is insufficient to account for geographic variation. Thus, the geographic variation results largely from genetic variation. Number of trunk vertebrae covaries with body size and shape in diverse vertebrate taxa, including Batrachoseps. I hypothesize that selection for different degrees of elongation, possibly related to fossoriality, has led to the extensive evolution of number of trunk vertebrae in Batrachoseps. Analysis of intrapopulational variation revealed sexual dimorphism in both body shape and number of trunk vertebrae, but no correlation between these variables in either sex. Females are more elongate than males, a pattern that has been attributed to fecundity selection in other taxa. Patterns of covariation among different classes of vertebrae suggest that some intrapopulational variation in number results from changes in vertebral identity rather than changes in segmentation.     

EVOLUTION OF PHENOTYPIC PLASTICITY IN THE STELLARIA LONGIPES COMPLEX: COMPARISONS AMONG CYTOTYPES AND HABITATS

Variation in the amount and pattern of plasticity was studied in three cytotypes (4x, 6x, and 8x) of Stellaria longipes and diploids of its suspected progenitor S. longifolia. All 13 traits considered showed plasticity. There were significant differences among cytotypes and habitats in plasticity for many traits. Overall, the diploids, S. longifolia, were most plastic, and the three cytotypes of S. longipes did not differ in amount of plasticity. Stellaria longifolia showed divergence from S. longipes in the pattern of plasticity as well. In general, cytotypes with more similar chromosome numbers had the same pattern of plasticity for more traits. Individuals from tundra populations differed in their pattern of plasticity from those of montane, boreal, and prairie origin, which were more similar to one another. Differences in plasticity among cytotypes were due primarily to divergence in amount, while differences among habitats were most often accounted for by divergent patterns of plasticity. We conclude that both polyploidy and natural selection have affected the evolution of plastic responses in this species complex. Analysis of the correlation between pairs of traits provided evidence that the pattern and amount of plasticity operate independently of one another and may be evolving separately.     

CONTRASTING PATTERNS OF PHENOTYPIC PLASTICITY IN REPRODUCTIVE TRAITS IN TWO GREAT TIT (PARUS MAJOR) POPULATIONS

Phenotypic plasticity is an important mechanism via which populations can respond to changing environmental conditions, but we know very little about how natural populations vary with respect to plasticity. Here we use random‐regression animal models to understand the multivariate phenotypic and genetic patterns of plasticity variation in two key life‐history traits, laying date and clutch size, using data from long‐term studies of great tits in The Netherlands (Hoge Veluwe [HV]) and UK (Wytham Woods [WW]). We show that, while population‐level responses of laying date and clutch size to temperature were similar in the two populations, between‐individual variation in plasticity differed markedly. Both populations showed significant variation in phenotypic plasticity (IxE) for laying date, but IxE was significantly higher in HV than in WW. There were no significant genotype‐by‐environment interactions (GxE) for laying date, yet differences in GxE were marginally nonsignificant between HV and WW. For clutch size, we only found significant IxE and GxE in WW but no significant difference between populations. From a multivariate perspective, plasticity in laying date was not correlated with plasticity in clutch size in either population. Our results suggest that generalizations about the form and cause of any response to changing environmental conditions across populations may be difficult.     

PHENOTYPIC PLASTICITY AND REACTION NORMS IN TEMPERATE AND TROPICAL POPULATIONS OF DROSOPHILA MELANOGASTER: OVARIAN SIZE AND DEVELOPMENTAL TEMPERATURE

The plasticity of ovariole number relative to developmental temperature was studied in three populations of Drosophila melanogaster at both ends of the cline: a temperate French population and two equatorial Congolese. Ovary size was much greater in the French flies, in agreement with an already known latitudinal cline. Among isofemale lines, significant differences in genetic variability were observed between populations with a maximum variability at intermediate temperatures. Parameters of phenotypic variability (CV and FA) were not statistically different among lines or populations, but a significant increase at low temperature was demonstrated for both. The shapes of the response curves (i.e., the norm of reaction) were analyzed by adjusting the data to a quadratic equation. The parameters of the equation were highly variable among lines. On the other hand, the temperature for maximum value of ovarioles (TMV) was much less variable and exhibited only a slightly significant difference between temperate and tropical flies (22.2°C vs. 22.7°C). During its geographic extension toward colder places, D. melanogaster underwent a large, presumably adaptative, increase in ovariole number but very little change in the norm of reaction of that trait.     

PHENOTYPIC PLASTICITY IN SCENEDESMUS COMMUNIS HEGEW. (CHLOROPHYCEAE). II. EXAMPLES OF ALGAL CYCLO- AND NONCYCLOMORPHOSIS

A new dimension to the well-known phenotypic plasticity in Scenedesmus is presented. Large colony type Scenedesmus communis Hegew. reproduced in a typical fashion in batch culture in standard media but produced small colonies (SCT) resembling S. komarekii Hegew. and S. subspicatus Chod. at low cell densities in dilute media. Highest frequency of production occurred after cells had been pretreated in a concentrated (inorganic nutrients) medium. Examples of both cyclic and noncyclic behavior in the life history of Scenedesmus are presented. The noncyclic behavior resembles previous reports on induced heritable changes in flax and tobacco due to different treatment applications of fertilizers. An analysis of gross morphological features using scanning electron microscopy showed that isolated strain SCT1 was similar to S. komarekii (strain UTEX 1236) and isolated strain SCT2 was similar to S. subspicatus (strain UTEX 1358). Growth and morphological expression of the induced SCT strains were highly similar to those of the respective UTEX species. The current state of taxonomy and the implications of phenotypic plasticity on taxonomy in Scenedesmus are discussed.     

INTRASPECIFIC LIFE HISTORY VARIATION IN A POND SNAIL: THE ROLES OF POPULATION DIVERGENCE AND PHENOTYPIC PLASTICITY

Two approaches were used to determine the degree of divergence in life histories among populations of the pond snail, Lymnaea elodes. Juvenile snails were reciprocally transferred between ponds differing in permanence and productivity, and the resulting variation in life history traits was recorded. In a second experiment, parents and their offspring from both a vernal and a permanent pond population were reared in the same pond. Proximal factors had by far the greatest effects on life history traits in the transfer experiment, with snails reared in a more productive pond showing earlier reproduction at a larger size, higher fecundity, and longer life cycle length. Snails from the more uncertain pond in terms of drying date did reproduce at an earlier age and smaller size and grew less in each pond. However, these population differences, for the most part, disappeared when snails were reared for two generations in the same environment. Much of the intraspecific variation in life histories seen in this species must therefore be considered the result of phenotypic plasticity. I argue that the plasticity in life histories itself may be adaptive to this inhabitant of unpredictable, vernal ponds.     

THE RESPONSES OF DROSOPHILA MELANOGASTER TO ARTIFICIAL SELECTION ON BODY WEIGHT AND ITS PHENOTYPIC PLASTICITY IN TWO LARVAL FOOD ENVIRONMENTS

To investigate the potential response to natural selection of reaction norms for age and size at maturity, fresh body weight at eclosion was mass selected under rich and poor larval food conditions in Drosophila melanogaster. The sensitivity of dry weight at eclosion to the difference between rich and poor larval food was selected using differences in sensitivities among families. For both experiments, the correlated response to selection of age at eclosion was examined. The flies were derived from wild populations and had been mass cultured in the lab for more than six months before the experiments started. These flies responded to selection on body weight upwards and downwards on both rich and poor larval food. Selection on increased or decreased sensitivity of body weight was also successful in at least one direction. Sensitivity was reduced by selection upwards in a poor environment and downwards in a rich environment.     

PHENOTYPIC PLASTICITY AND EPIGENETIC MARKING: AN ASSESSMENT OF EVIDENCE FOR GENETIC ACCOMMODATION

The relationship between genotype (which is inherited) and phenotype (the target of selection) is mediated by environmental inputs on gene expression, trait development, and phenotypic integration. Phenotypic plasticity or epigenetic modification might influence evolution in two general ways: (1) by stimulating evolutionary responses to environmental change via population persistence or by revealing cryptic genetic variation to selection, and (2) through the process of genetic accommodation, whereby natural selection acts to improve the form, regulation, and phenotypic integration of novel phenotypic variants. We provide an overview of models and mechanisms for how such evolutionary influences may be manifested both for plasticity and epigenetic marking. We point to promising avenues of research, identifying systems that can best be used to address the role of plasticity in evolution, as well as the need to apply our expanding knowledge of genetic and epigenetic mechanisms to our understanding of how genetic accommodation occurs in nature. Our review of a wide variety of studies finds widespread evidence for evolution by genetic accommodation.