Antipredator behavior promotes diversification of feeding strategies

Animals often facultatively engage in less risky behavior when predators are present. Few studies, however, have investigated whether, or how, such predator-mediated behavior promotes diversification. Here, we ask whether tadpoles of the spadefoot toad Scaphiopus couchii have a diminished ability to utilize a potentially valuable resource--anostracan fairy shrimp--because of behavioral responses to predation risk imposed by carnivorous tadpoles of the genus Spea. Observations of a congener of Sc. couchii that occurs in allopatry with Spea, coupled with an ancestral character state reconstruction, revealed that Sc. couchii's ancestors likely consumed shrimp. By experimentally manipulating the presence of Spea carnivore-morph tadpoles in microcosms, we found that Sc. couchii reduce feeding and avoid areas where both Spea carnivores and shrimp occur. We hypothesize that the recurrent expression of such behaviors in sympatric populations of Sc. couchii led to the evolutionary fixation of a detritivorous feeding strategy, which is associated with a reduced risk of predation from Spea carnivores. Generally, predator-mediated behavior might play a key role in promoting diversification of feeding strategies.     

ESS germination strategies in randomly varying environments. I. Logistic-type models

ESS germination strategies are studied in a model of annual plant population dynamics in a randomly varying environment. The possible strategies are different values of the annual germination fraction G, either constant over time or varying in response to a "cue" correlated with upcoming environmental conditions. The model generalizes D. Cohen's model (1966, J. Theor. Biol. 12, 119-129; 1968, J. Ecol. 56, 219-228) by allowing density-dependent per capita seed yields. ESSs are characterized in terms of the resulting harmonic mean growth rate of population density. The ESS criterion cannot be solved analytically, but qualitative relationships between the value of the ESS and other population parameters are obtained, and environments in which 100% germination is an ESS are identified. Some explicit predictions of the theory are summarized and compared with ideas of M. Westoby (1981, Amer. Nat. 118, 882-885). The results of this study are compared with those of Cohen (op. cit.) in a companion paper.     

盐生植物种子萌发对环境的适应对策

盐生环境是一种严峻的胁迫环境,对植物的生长、发育、繁殖等生活史的各阶段都产生着重要的影响。盐生植物是生长在盐渍土壤上的一类天然植物区系,它们在长期的进化过程中形成了一系列适应盐生生境的特殊生存策略。一般情况下,盐生植物种子对环境的适应能力,是植物对盐生环境适应性的重要体现;而植物发育早期对盐度的适应能力又是决定物种分布和群落组成的关键因素。在对国内外相关文献进行分析归纳的基础上,从盐分对种子萌发的影响机理及植物种子萌发对盐生环境的适应对策两个方面综述了植物种子休眠萌发与盐生环境的关系。     

Seed germination strategies of five perennial weeds

Summary The germination responses of seeds of Achillea millefolium L., Artemisia vulgaris L., Cirsium arvense (L.) Scop., Taraxacum officinale Weber, sensu lato, and Tussilago farfara L. to light, nitrate, alternating temperatures, chilling, light quality, and water availability were studied in laboratory tests, using fresh seed and seed stored for 6 months at 27°C and buried in the soil. A factorial experiment with light, nitrate, alternating temperatures, and seed age as factors found that all four affected germination except in T. farfara. All three external factors were stimulatory, especially in combinations. Fresh seed of A. vulgaris and C. arvense showed a light x alternating temperature synergism, responded to chilling, and after-ripened in cold dry storage. That of T. farfara had no dormancy and rapid germination, and germinated well on substrates with a water content too low for the other species. Seed of A. millefolium and A. vulgaris had good survival in both experimental storage conditions, while that of C. arvense and T. officinale did not, and that of T. farfara did not survive. Longevity in both conditions was associated with depth of initial dormancy. The two conditions caused different changes in dormancy in both A. vulgaris and A. millefolium. The germination behaviour, and the size, morphology and dispersal of the seeds of the species are discussed as strategies adapted to intermittently available situations for seedling establishment.     

ESS germination strategies in randomly varying environments. II. Reciprocal Yield-Law models

The study of ESS germination fractions in S. Ellner (1985, Theor. Pop. Biol. 27, 000-000) is applied and extended in the case of annuals obeying the Reciprocal Yield Law. The effects of parameter changes on the value of the ESS germination fraction are determined in some limiting cases by analyses of approximations. Numerical solutions of the ESS criterion are used to check the robustness of the conclusions and the accuracy of the approximations. In general, the ESS germination fraction decreases with an increased survivorship of buried seeds and with increased "variability" of seed yields. However, different measures of "variability" are appropriate in different circumstances. To avoid the possibility of conflicting predictions depending on the measure of variability, it is suggested that tests of the theory be limited to co-occurring species, and to variability due to climatic fluctuations. The ESS theory based on the Reciprocal Yield Law is compared with D. Cohen's (1966, J. Theor. Biol. 12, 119-129; 1968, J. Ecol. 56, 219-228) density-independent theory of "optimal" germination. The theories differ qualitatively and quantitatively regarding the influence of mean yield, seed survivorship, and the frequency of favorable years on the predicted germination fraction.     

Evolutionarily stable germination strategies with time-correlated yield

We investigate the effect of auto-correlated yield on the evolutionarily stable germination fraction of dormant seeds. By using both analytics and numerics, we first show that in a regime of small fluctuations a positive correlation reduces dormancy and a negative correlation enhances dormancy. By extending the numerical analysis we also show that in the regime of large fluctuations a more complex picture emerges where also negative correlations can reduce dormancy.     

Adaptive diversification of bitter taste receptor genes in Mammalian evolution

The diversity and evolution of bitter taste perception in mammals is not well understood. Recent discoveries of bitter taste receptor (T2R) genes provide an opportunity for a genetic approach to this question. We here report the identification of 10 and 30 putative T2R genes from the draft human and mouse genome sequences, respectively, in addition to the 23 and 6 previously known T2R genes from the two species. A phylogenetic analysis of the T2R genes suggests that they can be classified into three main groups, which are designated A, B, and C. Interestingly, while the one-to-one gene orthology between the human and mouse is common to group B and C genes, group A genes show a pattern of species- or lineage-specific duplication. It is possible that group B and C genes are necessary for detecting bitter tastants common to both humans and mice, whereas group A genes are used for species-specific bitter tastants. The analysis also reveals that phylogenetically closely related T2R genes are close in their chromosomal locations, demonstrating tandem gene duplication as the primary source of new T2Rs. For closely related paralogous genes, a rate of nonsynonymous nucleotide substitution significantly higher than the rate of synonymous substitution was observed in the extracellular regions of T2Rs, which are presumably involved in tastant-binding. This suggests the role of positive selection in the diversification of newly duplicated T2R genes. Because many natural poisonous substances are bitter, we conjecture that the mammalian T2R genes are under diversifying selection for the ability to recognize a diverse array of poisons that the organisms may encounter in exploring new habitats and diets.     

不同施肥处理对玉米秸秆碳氮比及其矿化特性的影响

以两个长期定位试验不同施肥处理玉米秸秆为对象,采用室内培养试验研究了其碳、氮养分在土壤中的矿化特性.结果表明：与未施肥处理相比,施用化肥（NPK）或化肥与有机肥配施（MNPK）处理明显增加了玉米秸秆的氮素含量,降低了其C/N.不同处理秸秆碳、氮矿化量和被微生物固持的碳、氮量因培养时期不同而异,NPK、MNPK和240 kg N·hm^-2处理秸秆在培养期间碳的矿化率显著高于相应不施肥处理;60 d培养期结束后,NPK处理秸秆的有机碳矿化量最大,占加入总有机碳的13.24%.各施肥处理玉米秸秆施入土壤后引起的土壤矿质氮固持量均较不施肥秸秆低,其中MNPK处理最低.施用秸秆增加了土壤微生物的代谢熵（qCO₂）,但不同处理间qCO₂的差异较小;各处理土壤微生物生物量碳、氮含量因培养时期不同而异.因此,生产中利用秸秆时应考虑不同施肥处理秸秆养分含量的差异.     

Rapid evolutionary divergence and ecotypic diversification of germination behavior in weedy rice populations

Feral plants have evolved from well-studied crops, providing good systems for elucidation of how weediness evolves. As yet, they have been largely neglected for this purpose. The evolution of weediness can occur by simple back mutations in domestication genes (domestication in reverse). Whether the evolutionary steps to weediness always occur in reverse remains largely unknown. We examined seed germination behavior in recently evolved weedy rice (Oryza sativa f. spontanea) populations and their coexisting cultivars in eastern and north-eastern China to address whether 'dedomestication' is the simple reverse of domestication. We found that these weedy populations did not diverge from their progenitors by reverting to the pre-domestication trait of seed dormancy. Instead, they have evolved a novel mechanism to avoid growing in inappropriate environments via changes in critical temperature cues for seed germination. Furthermore, we found evidence for subsequent ecotypic divergence of these populations such that the critical temperature for germination correlates with the local habitat temperature at latitudinal gradients. The origins of problematic plant species, weeds and invasives, have already been studied in detail. These plants can thus be used as systems for studying rapid evolution. To determine whether and how that evolution is adaptive, experiments such as those described here can be performed.     

Adaptive correlations between seed size and germination time

We present a model for the coevolution of seed size and germination time within a season when both affect the ability of the seedlings to compete for space. We show that even in the absence of a morphological or physiological constraint between the two traits, a correlation between seed size and germination time is nevertheless likely to evolve. This raises the more general question to what extent a correlation between any two traits should be considered as an a priori constraint or as an evolved means (or “instrument”) to actually implement a beneficial combination of traits. We derive sufficient conditions for the existence of a positive or a negative correlation. We develop a toy model for seed and seedling survival and seedling growth and use this to illustrate in practice how to determine correlations between seed size and germination time.

     

Adaptive changes in Plasmodium transmission strategies following chloroquine chemotherapy.

Both theory and data suggest that malaria parasites divert resources from within-host replication to the production of transmission stages (gametocytes) when conditions deteriorate. Increased investment into transmission stages should therefore follow subcurative treatment with antimalarial drugs, but relevant clinical studies necessarily lack adequate control groups. We therefore carried out controlled experiments to test this hypothesis, using a rodent malaria (Plasmodium chabaudi) model. Infections treated with a subcurative dose of the antimalarial chloroquine showed an earlier peak and a greater rate of gametocyte production relative to untreated controls. These alterations led to correlated changes in infectivity to mosquitoes, with the consequence that chloroquine treatment had no effect on the proportion of mosquitoes infected. Treatment of human malaria commonly does not result in complete parasite clearance. If surviving parasites produce compensatory increases in their rate of gametocyte production similar to those reported here, such treatment may have minimal effect on decreasing, and may actually increase, transmission. Importantly, if increased investment in transmission is a generalized stress response, the effect might be observed following a variety of antimalarial treatments, including other drugs and potential vaccines. Similar parasite life history counter-adaptations to intervention strategies are likely to occur in many disease-causing organisms.     

Adaptive diversification of a plastic trait in a predictably fluctuating environment

There is substantial evidence that evolutionary diversification can occur in allopatric conditions through reduction in the degree of phenotypic plasticity when an isolated population encounters a novel, more stable environment. Plasticity is no longer favored in the new environment, either because it carries an inherent physiological cost or because it leads to production of suboptimal phenotypes. In order to explore the role of phenotypic plasticity in sympatric diversification, we modeled the ecological and evolutionary dynamics of Escherichia coli bacteria in batch cultures. Our results describe an evolutionary pathway leading to metabolic diversification in a sympatric environment without spatial structure. In an environment that fluctuates widely and predictably, evolutionary branching leads to diversification and stable coexistence of generalist and specialist ecotypes for some combinations of parameters. Diversification and stable coexistence occur when reaction norms are steep and trade-offs between metabolic pathways are convex. We conclude that, in principle, diversification due to reduced plasticity can occur without allopatric isolation, reduced environmental variability, or an explicit cost of plasticity.     

Adaptive diversification in genes that regulate resource use in Escherichia coli

While there has been much recent focus on the ecological causes of adaptive diversification, we know less about the genetic nature of the trade-offs in resource use that create and maintain stable, diversified ecotypes. Here we show how a regulatory genetic change can contribute to sympatric diversification caused by differential resource use and maintained by negative frequency-dependent selection in Escherichia coli. During adaptation to sequential use of glucose and acetate, these bacteria differentiate into two ecotypes that differ in their growth profiles. The “slow-switcher” exhibits a long lag when switching to growth on acetate after depletion of glucose, whereas the “fast-switcher” exhibits a short switching lag. We show that the short switching time in the fast-switcher is associated with a failure to down-regulate potentially costly acetate metabolism during growth on glucose. While growing on glucose, the fast-switcher expresses malate synthase A (aceB), a critical gene for acetate metabolism that fails to be properly down-regulated because of a transposon insertion in one of its regulators. Swapping the mutant regulatory allele with the ancestral allele indicated that the transposon is in part responsible for the observed differentiation between ecological types. Our results provide a rare example of a mechanistic integration of diversifying processes at the genetic, physiological, and ecological levels.     

Adaptive strategies in natural populations of Drosophila

Summary Adult tolerance of ethanol vapour in a closed system containing 12% ethanol in solution, decreases in a cline from southern to northern Australia. However a Darwin population is more tolerant than predicted from its latitude. Ethanol tolerance races in Australia have almost certainly evolved within the last 100–150 years, because of resource unavailability prior to that time. Within populations, variation among isofemale strains is lowest in the climatically extreme southern Melbourne (37°S) and northern Darwin and Melville I. (11–12°S) populations. This suggests low resource diversity within extreme populations compared with the climatically less extreme Brisbane (28°S) and especially Townsville (19°S) populations. For desiccation resistance, the population rankings are: Darwin Melbourne > Townsville > Brisbane Melville I. and for development time, rankings are similar: Darwin Melbourne < Townsville < Brisbane Melville I.Therefore resource utilization heterogeneity is greatest in populations not greatly stressed by desiccation and where development times are extended. In total therefore, the utilization of a diversity of resources is a feature of populations tending somewhat towards a K-strategy; this is emphasized by the relative heterogeneity among isofemale strains of these populations for desiccation resistance and to a lesser extent development times.The D. melanogaster gene pool can be viewed as made up of climate-associated races. Since the ethanol tolerances of adjacent (and climatically similar) Darwin and Melville I. are very different, resource utilization races may occur within climatic races. Such a mosaic of resource utilization races are more likely in climatically extreme than in optimal habitats.     

Rapid diversification of male genitalia and mating strategies in Ohomopterus ground beetles

We analysed evolutionary diversification and covariation in male genitalia and four mating traits related to sexual selection, i.e. testis size, spermatophore size, copulation duration and post-copulatory guarding duration, in Ohomopterus ground beetles using phylogenetically independent contrasts. Male genital size and mating duration have evolved more rapidly than body size and the other traits studied. Male genital size was negatively correlated with copulation duration, suggesting that elongated male genitalia may enable decreased time investment in a single copulation because it is more effective at facilitating spermatophore deposition. Male genital size was positively correlated with spermatophore size, suggesting coevolution between offensive and defensive male mating tactics because the elongated male genitalia may be advantageous in displacement of rivals' plug-like spermatophores, and decreased mating duration may intensify sperm competition. Thus, the remarkable diversity of male genitalia in Ohomopterus may have been facilitated by the interplay between inter- and intrasexual selection processes.     

Seed dormancy and germination of five selected NATURA-2000 plant species from Croatia showing different germination strategies

Abstract

NATURA-2000, the ecological network of protected areas in the European Union that has been included in the Croatian legislation, defines Community important plant species with imperative on their conservation ex/in situ. In the Botanical Garden of the Faculty of Science, University of Zagreb, five NATURA-species have been selected as research subjects for germination study to shed light on the topic of their seed ecology and consequently advance their conservation efforts: Degenia velebitica, Scilla litardierei, Klasea lycopifolia, Ligularia sibirica and Genista holopetala. The freshly matured seeds of each species were exposed to cold or warm stratification in duration of four to sixteen weeks, and their germination was investigated through different regimes of incubation parameters, i.e. illumination (light/dark) and temperature (5, 15/6, 23?°C). All species had higher germination values after cold stratification, with the exception of G. holopetala. We concluded that D. velebitica, K. lycopifolia and presumably L. sibirica seeds has non-deep physiological dormancy while S. litardierei has deep complex morphophysiological dormancy and G. holopetala has physical dormancy. The observed patterns in seeds’ behaviour are consistent with the conditions in their natural habitats in Croatia and the knowledge of these patterns is vital for successful conservation strategies in the future.