QUANTITATIVE GENETICS OF BRYOZOAN PHENOTYPIC EVOLUTION. III. PHENOTYPIC PLASTICITY AND THE MAINTENANCE OF GENETIC VARIATION

Cheilostome bryozoan species show long-term morphologic stasis, implying stabilizing selection sustained for millions of years, but nevertheless retain significant heritable variation in traits of skeletal morphology. The possible role of within-genotype (within-colony) phenotypic variability in preserving genetic diversity was analyzed using breeding data for two species of Stylopoma from sites along 110 km of the Caribbean coast of Panama. Variation among zooids within colonies accounts for nearly two-thirds of the phenotypic variance on average, increases with environmental heterogeneity, and includes significant genotype-environment interaction. Thus, within-colony variability apparently represents phenotypic plasticity, at least some of which is heritable, rather than random “developmental noise.” Almost all of the among-colonies component of phenotypic variance is accounted for by additive genetic differences in trait means, suggesting that within-colony plasticity includes virtually all of the environmental component of phenotypic variance in these populations of Stylopoma. Thus, heritable within-colony plasticity could play a significant part in maintaining genetic diversity in cheilostomes, but it is also possible that rates of polygenic mutation alone are sufficient to balance the effects of selection.     

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.     

POPULATION DIFFERENTIATION FOR PHENOTYPIC PLASTICITY IN THE STELLARIA LONGIPES COMPLEX

Population differentiation for phenotypic plasticity of 12 morphological and reproductive traits was investigated in five populations of the Stellaria longipes complex including a population of the sand dune endemic S. arenicola. Population differentiation was detected for the mean (genotypic) value, amount of plasticity, and pattern of plasticity of traits. Average amount of plasticity was not related to degree of isozyme variability in the populations. Differentiation for pattern of plasticity was much more common than for amount. The direction and extent of divergence among populations was dependent on which of the three trait aspects was under consideration (mean, amount of plasticity, pattern of plasticity) and did not reflect their similarity as revealed by enzyme electrophoretic data. It was concluded that trait means, amounts of plasticity, and patterns of plasticity are independent of one another during evolutionary divergence and may be influenced by mosaic selection.     

GENOTYPE-ENVIRONMENT INTERACTION AND THE EVOLUTION OF PHENOTYPIC PLASTICITY

Studies of spatial variation in the environment have primarily focused on how genetic variation can be maintained. Many one-locus genetic models have addressed this issue, but, for several reasons, these models are not directly applicable to quantitative (polygenic) traits. One reason is that for continuously varying characters, the evolution of the mean phenotype expressed in different environments (the norm of reaction) is also of interest. Our quantitative genetic models describe the evolution of phenotypic response to the environment, also known as phenotypic plasticity (Gause, 1947), and illustrate how the norm of reaction (Schmalhausen, 1949) can be shaped by selection. These models utilize the statistical relationship which exists between genotype-environment interaction and genetic correlation to describe evolution of the mean phenotype under soft and hard selection in coarse-grained environments. Just as genetic correlations among characters within a single environment can constrain the response to simultaneous selection, so can a genetic correlation between states of a character which are expressed in two environments. Unless the genetic correlation across environments is ± 1, polygenic variation is exhausted, or there is a cost to plasticity, panmictic populations under a bivariate fitness function will eventually attain the optimum mean phenotype for a given character in each environment. However, very high positive or negative correlations can substantially slow the rate of evolution and may produce temporary maladaptation in one environment before the optimum joint phenotype is finally attained. Evolutionary trajectories under hard and soft selection can differ: in hard selection, the environments with the highest initial mean fitness contribute most individuals to the mating pool. In both hard and soft selection, evolution toward the optimum in a rare environment is much slower than it is in a common one. A subdivided population model reveals that migration restriction can facilitate local adaptation. However, unless there is no migration or one of the special cases discussed for panmictic populations holds, no geographical variation in the norm of reaction will be maintained at equilibrium. Implications of these results for the interpretation of spatial patterns of phenotypic variation in natural populations are discussed.     

NATURAL HISTORY AND POPULATION DYNAMICS OF THE LAND SNAIL HELIX TEXTA IN ISRAEL (PULMONATA: HELICIDAE)

Helix texta is endemic to the Mediterranean regions of Israel.It has a seasonal activity pattern which starts in the autumn,with the first rains, and dwindles towards the spring, whenthe snails dig into the ground for a six months long aestivation.A cold spell of 0°C will, however, terminate the activeseason of the adult snail, even in the middle of the rainy season.Survival of the young is very low and most of them (90%) donot survive their first year, because of the winter cold andthe summer drought. Massive predation of adult snails by wild boar was observedin December 1986: within a few days, about 50% of the adultsin the study in the area were eaten. A very rapid growth ofyoung and subadults was observed immediately after this predation.These observations suggest that the extent of recruitment ofnew adults to the population may be partly controlled by existingadults, through a growth-inhibiting pheromone in the mucus.After predation this inhibiting factor disappears, enablinga subsequent rapid growth of the young. In this manner, massive,irregular predation of the adults by a large predator, and changesin juvenile survival, result in sharp fluctuations in the agestructure of the population. The resulting pattern of unstablepopulation dynamics is different from that described for theEuropean species of Helix. (Received 16 January 1989; accepted 17 April 1989)     

MICROGEOGRAPHIC GENETIC STRUCTURE OF MORPHOLOGICAL AND LIFE HISTORY TRAITS IN A NATURAL POPULATION OF IMPATIENS CAPENSIS

To investigate the microgeographic spatial structure of genetic variation for quantitative traits in a natural population of Impatiens capensis, we performed a common-garden greenhouse experiment. Seedlings were collected at 10-m intervals from a 40 times 40-m permanent grid in a natural population and grown to maturity in a greenhouse. From these parents, 3 self-fertilized seed families per grid point were then grown in a randomized design in the greenhouse and scored for a variety of morphological and life-history traits. Virtually all of the traits displayed significant variation among families, and many were significantly heterogeneous among grid points, indicating microgeographic genetic differentiation on a fine spatial scale. Overall morphological divergence, measured as Mahalanobis distances between grid points, increased with geographical distance. In general, spatial autocorrelation coefficients of grid point character means were positive at 11–20 m and negative beyond 40 m, although power for significance testing was low. The first factor in a principal component analysis of grid point means was positively loaded on height-related traits and negatively loaded on total reproduction at 50 days, accounting for 31% of the variation. This factor displayed significant positive spatial autocorrelation at 11–20 m and negative autocorrelation at >40 m. The remaining factors showed no detectable spatial structuring among grid points. These differences in spatial pattern among characters suggest that forces other than drift may have influenced the genetic structure of the population. There was no evidence for density-dependent selection; seedling density was not significantly correlated with the grid point mean of any trait.     

GEOGRAPHIC VARIATION IN THE LIFE HISTORY OF MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Variation in the geographic distribution of the life histories of Mastocarpus papillatus was investigated. Carpospores were isolated from 377 female gametophytes collected from eight localities on the Pacific coast of Baja California, Mexico, and California, U.S.A., and grown in laboratory culture. All carpospores from a single female gave rise either to basal discs with gametophyte-like uprights or crustose plants formerly referred to the genus Petrocelis. Early stages in the development of each type of germling were observed, and environmental factors affecting development were suggested. Based on carpospore germlings, females from each location were scored as having either the 1) sexual life history (crustose germlings) or 2) direct-development life history (discoid germlings with uprights). All females from the two southernmost locations in Baja California exhibited the sexual life history. In the three locations from the central-southern California coast, 70-95% of the females exhibited the sexual life history and the remainder exhibited the direct-development life history. In two of the three populations from the central-northern California coast, 70-90% of the females exhibited the direct-development life history and the remainder the sexual life history. In the third location from the central-northern California coast, the northernmost location sampled in the current study, 60% of the females exhibited the sexual life history and 40% the direct-development life history. The relative ecological advantages and disadvantages of the life histories are unknown as are the environmental factors that produced the ratios of sexual to direct-development females observed at each location.     

PLASTICITY OF THE LIFE CYCLE OF XEROPICTA DERBENTINA (KRYNICKI, 1836), A RECENTLY INTRODUCED SNAIL IN MEDITERRANEAN FRANCE

Xeropicta derbentina (Krynicki, 1836), a native of Eastern MediterraneanEurope, was introduced to southeastern France during the 1940sand is now widely spread across Provence. In summer it aggregateson plants, making its populations clearly visible. However,its life cycle within the Mediterranean basin is poorly documented.While X. derbentina in its native area exhibits an annual lifecycle, this species has been found in Provence to have a bienniallife cycle. Moreover, in southeastern France, field studieswithin a restricted area show variations in demographic structure.In consequence, the life cycle of X. derbentina and the demographicpatterns observed require clarification. Five populations withvarious demographic structures were studied over 1 year in thesame location, i.e. under the same climatic conditions. Thefield study was complemented by laboratory observations on mating,egg-laying and hatching. Xeropicta derbentina appears to bea semelparous species, with an annual life cycle being foundon four plots. The reproductive period begins at the end ofsummer and lasts until the beginning of winter. First egg-layingoccurs within 1 week after mating and lasts up to 30 days. Hatchingtakes place 15–20 days after egg-laying. Xeropicta derbentinapossesses multiple mating and egg-laying sessions, involvingsuccessive hatching. Populations are mainly characterized bytwo growth stages, the first in spring when newly-hatched snailsevolve into juveniles, and the second in late summer when theyreach maturity. However, on the highest density plot, a bienniallife cycle is observed for some newly-hatched snails that showan interrupted growth during summer and evolve into juvenilesonly in the second autumn. Moreover, this life cycle not onlyvaries among plots but also at a 1-year interval within plots.Hence, the life span of X. derbentina is between 12 and 20 months,but can be extended up to 30 months according to whether hatchingoccurs early or late and whether they survive the first andsecond winters. Xeropicta derbentina is thus able to have variousgrowth speeds and life spans, and appears to switch from anannual life cycle to a biennial cycle in response to populationdensity or climatic conditions. (Received 8 October 2004; accepted 15 December 2004)     

INTRASPECIFIC VARIATION IN ANT SEX RATIOS AND THE TRIVERS-HARE HYPOTHESIS

We consider worker-controlled sex investments in eusocial Hymenoptera (ants in particular) and assume that relatedness asymmetry is variable among colonies and that workers are able to assess the relatedness asymmetry in their own colony. We predict that such “assessing” workers should maximize their inclusive fitness by specializing in the production of the sex to which they are relatively most related, i.e., colonies whose workers have a relatedness asymmetry below the population average should specialize in males, whereas colonies whose workers have a higher than average relatedness asymmetry should specialize in making females. Our argument yields the expectation that colony sex ratios will be bimodally distributed in ant populations where relatedness asymmetry is variable owing to multiple mating, worker reproduction, and/or polygyny. No such bimodality is expected, however, in ant species where relatedness asymmetry is known to be constant, or in cases where relatedness asymmetry is supposed to be irrelevant due to allospecific brood rearing under queen control, as in the slave-making ants. Comparative data on colony sex ratios in ants are reviewed to test the predictions. The data partly support our contentions, but are as yet insufficient to be considered as decisive evidence.     

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.