INBREEDING DEPRESSION UNDER JOINT SELFING,OUTCROSSING, AND ASEXUALITY

Partial asexual reproduction was introduced into a model of inbreeding depression due to nearly recessive lethal mutations in a partially selfing population. The frequencies of asexuality, selfing, and outcrossing were either constant or occurred in cycles of a single sexual generation followed by one or more asexual generations. We found that increasing the degree of asexuality generally increases the inbreeding depression maintained in an equilibrium population with a given selfing rate. This is due to the increase in the number of mutations relative to sexual generations during which selfing-induced purging of mutations may take place. For very high genomic mutation rates, sufficient to produce a threshold rate of self-fertilization for purging recessive lethal mutations, asexuality can have the opposite effect, decreasing equilibrium inbreeding depression, because of an increase in the efficiency of selection against mutations in heterozygotes with asexuality.     

An analysis of the growth of the retinal cell population in embryonic chicks yielding proliferative ratios, numbers of proliferative and non-proliferative cells and cell-cycle times for successive generations of cell cycles

Growth curves of the retinal cell population of embryonic chicks were fitted by a branching-process model of cell population growth, thereby estimating the proliferative ratios and mean cell-cycle times of the generations of cell cycles that underlie retinal growth. The proliferative ratio determines the proportion of cells that divides in the next generation, so the numbers of proliferative and non-proliferative cells in each generation of cell cycles were obtained. The mean cell-cycle times determine the times over which the generations are extant. Assuming growth starts from one cell in generation 0, the proliferative cells reach 3.6 × 10⁶ and the non-proliferative cells reach 1.1 × 10⁶ by generation 23. The next four generations increase the proliferative cell numbers to 13.9 × 10⁶ and produce 20.1 × 10⁶ non-proliferative cells. In the next five generations in the end phase of growth, non-proliferative cells are produced in large numbers at an average of 13.9 × 10⁶ cells per generation as the retinal lineages are completed. The retinal cell population reaches a maximum estimated here at 98.2 × 10⁶ cells. The mean cell-cycle time estimates range between 6.8 and 10.1 h in generations before the end phase of growth and between 10.6 and 17.2 h in generations in the end phase. The retinal cell population growth is limited by the depletion of the proliferative cell population that the production of non-proliferative cells entails. The proliferative ratios and the cell-cycle-time distribution parameters are the likely determinants of retinal growth rates. The results are discussed in relation to other results of spatial and temporal patterns of the cessation of cell cycling in the embryonic chick retina.     

Stable periodic solutions for two species,density dependent coevolution

This paper studies a four dimensional system of time-autonomous ordinary differential equations which models the interaction of two diploid, diallelic populations with overlapping generations. The variables are two population densities and an allele frequency in each of the populations. For single species models, the existence of periodic solutions requires that the genotype fitness functions be both frequency and density dependent. But, for two species exhibiting a predator-prey interaction, two examples are presented where there exists asymptotically stable cycles with fitness functions only density dependent. In the first example, the Hopf bifurcation theorem is used on a two parameter, polynomial vector field. The second example has a Michaelis-Menten or Holling term for the interaction between predator and prey; and, for this example, the existence and uniqueness of limit cycles for a wide range of parameter values has been established in the literature.     

Developmental synchrony in multivoltine insects: generation separation versus smearing

Many insect species undergo multiple generations each year. They are found across biomes that vary in their strength of seasonality and, depending on location and species, can display a wide range of population dynamics. Some species exhibit cycles with distinct generations (developmental synchrony/generation separation), some exhibit overlapping generations with multiple life stages present simultaneously (generation smearing), while others have intermediate dynamics with early season separation followed by late season smearing. There are two main hypotheses to explain these dynamics. The first is the ‘seasonal disturbance’ hypothesis where winter synchronizes the developmental clock among individuals, which causes transient generation separation early in the season that erodes through the summer. The second is the ‘temperature destabilization’ hypothesis where warm temperatures during the summer cause population dynamics to become unstable giving rise to single generation cycles. Both hypotheses are supported by detailed mathematical theory incorporating mechanisms that are likely to drive dynamics in nature. In this review, we synthesize the theory and propose a conceptual framework—where each mechanism may be seen as an independent axis shaping the developmental (a)synchrony—that allows us to predict dynamic patterns from insect life-history characteristics. High fecundity, short adult life-span and strong seasonality enhance synchrony, while developmental plasticity and environmental heterogeneity erode synchrony. We further review current mathematical and statistical tools to study multi-generational dynamics and illustrate using case studies of multivoltine tortrix moths. By integrating two disparate bodies of theory, we articulate a deep connection among temperature, stability, developmental synchrony and inter-generational dynamics of multivoltine insects that is missing in current literature.     

Spontaneous and Ethyl Methanesulfonate-Induced Mutations Controlling Viability in DROSOPHILA MELANOGASTER. II. Homozygous Effect of Polygenic Mutations

Polygenic mutations affecting viability were accumulated on the second chromosome of Drosophila melanogaster by treating flies with EMS in successive generations. The treated chromosomes were later made homozygous and tested for their effects on viability by comparison of the frequency of such homozygotes with that of other genotypes in the same culture. The treated wild-type chromosomes were kept heterozygous in Pm/+ males by mating individual males in successive generations to Cy/Pm females. The number of generations of accumulation was 1 to 30 generations, depending on the concentration of EMS. A similar experiment for spontaneous polygenic mutations was also conducted by accumulating mutations for 40 generations. The lower limit of the spontaneous mutation rate of viability polygenes is estimated to be 0.06 per second chromosome per generation, which is about 12 times as high as the spontaneous recessive lethal mutation rate, 0.005. EMS-induced polygenic mutations increase linearly with the number of treated generations and with the concentration of EMS. The minimum mutation rate of viability polygenes is about 0.017 per 10(-4)m, which is only slightly larger than the lethal rate of 0.013 per 10(-4) m. The maximum estimate of the viability reduction of a single mutant is about 6 to 10 percent of the normal viability. The data are consistent with a constant average effect per mutant at all concentrations, but this is about three times as high as that for spontaneous mutants. It is obvious that one can obtain only a lower limit for the mutation rate, since some mutants may have effects so near to zero that they cannot be detected. The possibility of measuring something other than the lower limit is discussed. The ratio of the load due to detrimental mutants to that caused by lethals, the D/L ratio, is about 0.2 to 0.3 for EMS-induced mutants, as compared to about 0.5 for spontaneous mutants. This is to be expected if EMS treatment produces a large fraction of small deletions and other chromosome rearrangements which are more likely to be lethal.     

An in vitro whole plant selection system: paraquat tolerant mutants in the fern Ceratopteris

Summary A whole plant selection system using the haploid gametophyte generation of the fern Ceratopteris richardii has been developed to select for mutations that confer resistance or tolerance to various selection pressures. The expression of the mutations can be analyzed and characterized in both the haploid gametophyte and diploid sporophyte generations. Genetic analyses are facilitated by the fern's rapid life cycle and the ease of manipulating the gametophyte generation. Selection for tolerance to the herbicide paraquat has yielded two mutants which have an increased tolerance to the herbicide in both the gametophyte and sporophyte generations. Both mutants exhibit single nuclear gene inheritance patterns and appear to be closely linked or allelic.     

Measuring selection in human populations using the growth rate per generation

Estimates of the speed of evolution between generations depend on the association between individual traits and a measure of fitness. The two most frequently used measures of fitness are the net reproduction rate and the 1-year growth factor implied by the fertility and mortality rates. Results based on the two lead to very different results. The reason is that the 1-year growth factor is not a measure of change between generations. Therefore, studies of changes between generations should use the amount of growth over the length of a generation. This is especially important for studies of human populations because of the long length of generation. In addition, estimates based on a single year''s growth are overly sensitive to data on individuals who fail to reproduce. The effects of using a generational measure are demonstrated using data from Kenya and Ukraine. These results demonstrate that using a 1-year growth rate to measure fitness leads to estimates that understate the rate at which evolution changes the characteristics of a human population.     

On the formula for admixture linkage disequilibrium

Admixture linkage disequilibrium (ALD), a phenomenon created by gene flow between genetically distinct populations, has for some time been used as a tool in gene mapping. It is therefore important to analyze the pattern of ALD over generations. In this study we explore two models of admixture: the gradual admixture (GA) model, in which admixture occurs at a variable rate in every generation; and the immediate admixture (IA) model, a special case of the GA model, in which admixture occurs in a single generation. In the case of ALD, the well-known formula of linkage disequilibrium (Delta(t)=(1-r)t Delta(0)) is not applicable under these two models. We note the effect of a random mating population (RMP) to the gametic frequencies from the parental population to the offspring population, and provide the correct formula for ALD.     

Contrasting gene expression programs correspond with predator‐induced phenotypic plasticity within and across generations in Daphnia

Research has shown that a change in environmental conditions can alter the expression of traits during development (i.e., “within‐generation phenotypic plasticity”) as well as induce heritable phenotypic responses that persist for multiple generations (i.e., “transgenerational plasticity”, TGP). It has long been assumed that shifts in gene expression are tightly linked to observed trait responses at the phenotypic level. Yet, the manner in which organisms couple within‐ and TGP at the molecular level is unclear. Here we tested the influence of fish predator chemical cues on patterns of gene expression within‐ and across generations using a clone of Daphnia ambigua that is known to exhibit strong TGP but weak within‐generation plasticity. Daphnia were reared in the presence of predator cues in generation 1, and shifts in gene expression were tracked across two additional asexual experimental generations that lacked exposure to predator cues. Initial exposure to predator cues in generation 1 was linked to ~50 responsive genes, but such shifts were 3–4× larger in later generations. Differentially expressed genes included those involved in reproduction, exoskeleton structure and digestion; major shifts in expression of genes encoding ribosomal proteins were also identified. Furthermore, shifts within the first‐generation and transgenerational shifts in gene expression were largely distinct in terms of the genes that were differentially expressed. Such results argue that the gene expression programmes involved in within‐ vs. transgeneration plasticity are fundamentally different. Our study provides new key insights into the plasticity of gene expression and how it relates to phenotypic plasticity in nature.     

Fitness of Virulent Meloidogyne incognita Isolates on Susceptible and Resistant Cowpea

A study of life-history traits was made to determine factors associated with the fitness of Meloidogyne incognita isolates virulent to resistance gene Rk in cowpea. Egg hatch, root penetration, egg mass production, and fecundity (eggs per egg mass) of avirulent and virulent phenotypes were compared among M. incognita isolates, isofemale lines, and single descent lines over multiple generations on resistant and susceptible cowpea. Variation (P ≤ 0.05) in both hatch and root penetration rates was found among isolates at a given generation. However, this variation was not consistent within nematode lines among generations, and there was no correlation with level of virulence, except for penetration and virulence on resistant cowpea at generation 20. Resistant and susceptible cowpea roots were penetrated at similar levels. Differences in reproductive factors on resistant plants were correlated with levels of virulence expression. In some isofemale lines, single descent lines, and isolates, lower (P ≤ 0.05) rates of egg mass production and fecundity on susceptible cowpea were associated with virulence to Rk, indicating a trade-off between reproductive fitness and virulence. Other virulent nematode lines from the same isolates did not have reduced reproductive ability on susceptible cowpea over 27 generations. Thus, virulent lineages varied in reproductive ability on susceptible cowpea, contributing to adaptation and maintenance of virulence within M. incognita populations under stabilizing selection.     

马尾松毛虫2、3代分化和干旱对种群时间动态的影响

通过对安徽冬至县金寺山林场27年中马尾松毛虫Dendrolimus punctatus Walker发生面积的时间序列和2、3代分化率及干旱记录进行相关分析,进一步探讨2、3代分化和干旱对马尾松毛虫种群时间序列动态的影响。马尾松毛虫2、3代分化率有一定的周期性,而且波动的幅度很大,大致可以分成低、中、高三个等级,三个等级的分化率范围分别为1%～10%,11%～40%和41%～100%。大致每隔5～6年出现一次高分化率,高分化率的前后往往是低分化率,高分化率之间一般是中等的分化率和低分化率交替出现。通过分析马尾松毛虫2、3代分化对种群时间序列动态的影响,说明2、3代分化主要影响当代的数量,与以后各代数量无明显相关。2、3代分化前的种群数量与2、3代分化也无明显的相关。中等以上的分化率常引起松毛虫的发生。影响2、3代分化的主要气候因子是7月份的气温,7月份的气温越高,卵孵化的高峰期越早,更多幼虫于分化的临界光周期以前孵化,分化率就高。干旱与2、3代分化有微弱的相关,对松毛虫时间动态的影响具有延迟作用,影响最明显的是第二年的第1代,这说明干旱不仅影响当代松毛虫的动态,更重要的是通过影响寄主,从而影响到以后世代的松毛虫动态。     

Life-cycle-generation-specific developmental processes are modified in the immediate upright mutant of the brown alga Ectocarpus siliculosus

Development of the sporophyte and gametophyte generations of the brown alga E. siliculosus involves two different patterns of early development, which begin with either a symmetric or an asymmetric division of the initial cell, respectively. A mutant, immediate upright (imm), was isolated that exhibited several characteristics typical of the gametophyte during the early development of the sporophyte generation. Genetic analyses showed that imm is a recessive, single-locus Mendelian factor and analysis of gene expression in this mutant indicated that the regulation of a number of life-cycle-regulated genes is specifically modified in imm mutant sporophytes. Thus, IMM appears to be a regulatory locus that controls part of the sporophyte-specific developmental programme, the mutant exhibiting partial homeotic conversion of the sporophyte into the gametophyte, a phenomenon that has not been described previously.     

A multigenerational effect of parental age on offspring size but not fitness in common duckweed (Lemna minor)

Classic theories on the evolution of senescence make the simplifying assumption that all offspring are of equal quality, so that demographic senescence only manifests through declining rates of survival or fecundity. However, there is now evidence that, in addition to declining rates of survival and fecundity, many organisms are subject to age‐related declines in the quality of offspring produced (i.e. parental age effects). Recent modelling approaches allow for the incorporation of parental age effects into classic demographic analyses, assuming that such effects are limited to a single generation. Does this ‘single‐generation’ assumption hold? To find out, we conducted a laboratory study with the aquatic plant Lemna minor, a species for which parental age effects have been demonstrated previously. We compared the size and fitness of 423 laboratory‐cultured plants (asexually derived ramets) representing various birth orders, and ancestral ‘birth‐order genealogies’. We found that offspring size and fitness both declined with increasing ‘immediate’ birth order (i.e. birth order with respect to the immediate parent), but only offspring size was affected by ancestral birth order. Thus, the assumption that parental age effects on offspring fitness are limited to a single generation does in fact hold for L. minor. This result will guide theorists aiming to refine and generalize modelling approaches that incorporate parental age effects into evolutionary theory on senescence.     

Principles of the organization of intestinal epithelium and its recovery from damage revealed by modeling

Using the developed method of modelling the curves of the fraction of labelled indexes (FLI) in the small intestine epithelium, a relationship was found between the FLI shape and duration of the S phase and the whole generation cycle for various generations of proliferating crypt cells. On the basis of the comparison between the theoretical and experimental FLI the normal values of the generation cycles along the crypt were specified for various intestinal parts. FLI were shaped along the crypt at various time intervals after irradiation. As a result, the data were obtained on the correlation between the duration of generation cycles and their phases during the postirradiation period and for various cell generations.     

Layered distribution, according to age, within the cell wall of bacillus subtilis.

When soluble autolytic activity was added to growing cultures of a mutant possessing a reduced rate of cell wall turnover, there was a delay of more than one generation before solubilization of new cell wall began, in contrast to the immediate increase in the rate of solubilization of old cell wall. A similar delay was found before turnover of new cell wall occurred in the parent, in agreement with a previous report (Mauck et al., 1971). When sodium lauryl sulfate-inactivated cell walls were prepared, the great bulk of the wall formed a uniformly susceptible substrate to added autolytic activity. The immediate solubilization of new wall eliminates insusceptibility to autolytic enzyme as an explanation for the failure to be turned over. There were, however, major differences in the rate of solubilization of wall of different ages. During solubilization of the initial 30% of the cell wall preparation, wall two generations old was solubilized at least seven times faster than wall one-half a generation old. This result is interpreted in terms of differences in accessibility. The cell wall is seen as consisting of a series of layers, the age of which increases with the distance from the membrane, such that wall newly synthesized on the membrane passes out through the thickness of the cell wall layer during subsequent growth and only becomes susceptible to turnover as it reaches the outer surface, largely in the form of a layer, more than one generation after incorporation.     

Population dynamical consequences of gregariousness in a size-structured consumer-resource interaction

Many animal species live in groups. Group living may increase exploitation competition within the group, and variation among groups in intra-group competition intensity could induce life-history variability among groups. Models of physiologically structured populations generally predict single generation cycles, driven by exploitation competition within and between generations. We expect that life-history variability and habitat heterogeneity induced by group living may affect such competition-driven population dynamics. In this study, we vary the gregariousness (the tendency to aggregate in groups) of a size-structured consumer population in a spatially explicit environment. The consumer has limited mobility, and moves according to a probabilistic movement process. We study the effects on the population dynamics, as mediated through the resource and the life-history of the consumer. We find that high gregariousness leads to large spatial resource variation, and highly variable individual life-history, resulting in highly stochastic population dynamics. At reduced gregariousness, life-history of consumers synchronizes, habitat heterogeneity is reduced, and single generation cycles appear. We expect this pattern to occur for any group living organism with limited mobility. Our results indicate that constraints set by population dynamical feedback may be an important aspect in understanding group living in nature.     

FERTILIZATION SUCCESS CAN DRIVE PATTERNS OF PHASE DOMINANCE IN COMPLEX LIFE HISTORIES1

Many algal life cycles alternate between two free‐living generations. Life histories in which the two generations look identical (isomorphic) are common, particularly in the Rhodophyta. Reports of natural populations dominated by one generation of the life history have sought explanation in terms of phase‐specific differences in mortality and reproductive output, yet in many cases identification of these adaptations has been elusive or inconsistent with predictions. We hypothesized that the gametophyte‐to‐sporophyte ratio of ecologically equivalent isomorphs could result from variation in fertilization rate. We developed two models to test this hypothesis: one representing a generalized isomorphic life history and the other specific to red algae with a Polysiphonia‐type life history. Fertilization rate affected the gametophyte‐to‐sporophyte ratio, especially at low fertilization rates. In the general model, gametophytes dominated the population regardless of fertilization rate unless egg production greatly exceeded meiospore production. In the red algal model, phase dominance depended on the combination of fertilization rate and the number of carpospores produced per fertilization. The generational composition of model multiphasic algal populations results from their inherent reproductive characteristics and the dynamic environment to which fertilization and mortality rates are tied.     

Analysis of Inheritability and Expression Profile of Single and Multi-copy Transgene(s) in Rice Over Generations

Transformation of Oryza sativa subsp indica variety Pusa Basmati 1 with Agrobacterium tumefaciens strain LBA4404(pTOK233) carrying genes coding for neomycin phosphotransferase (nptII), β-glucuronidase (gus) and hygromycin phosphotransferase (hph) under the control of plant-specific promoters (pnos and pCaMV35S) within its T-DNA region produced transgenics with single and multiple copies of T-DNA. Simple Mendelian as well as complex patterns of the inheritance for hygromycin resistance trait were observed in R1 and R2 generations. Non-segregating lines selected in R2 generation did not show further segregation of the resistance trait in R3 and R4 generations accompanied by stabilization of integrated transgenes. One of these lines showed the presence of truncated T-DNA in R1 generation. The single copy transgenics showed high stability of expression of gus gene, whereas multi-copy transgenics were prone to silencing up to R3 generation after which no further reduction in gene expression was observed.     

Imprinting capacity of gamete lineages in Caenorhabditis elegans

We have observed a gamete-of-origin imprinting effect in C. elegans using a set of GFP reporter transgenes. From a single progenitor line carrying an extrachromosomal unc-54::gfp transgene array, we generated three independent autosomal integrations of the unc-54::gfp transgene. The progenitor line, two of its three integrated derivatives, and a nonrelated unc-119:gfp transgene exhibit an imprinting effect: single-generation transmission of these transgenes through the male germline results in approximately 1.5- to 2.0-fold greater expression than transmission through the female germline. There is a detectable resetting of the imprint after passage through the opposite germline for a single generation, indicating that the imprinted status of the transgenes is reversible. In cases where the transgene is maintained in either the oocyte lineage or sperm lineage for multiple, consecutive generations, a full reset requires passage through the opposite germline for several generations. Taken together, our results indicate that C. elegans has the ability to imprint chromosomes and that differences in the cell and/or molecular biology of oogenesis and spermatogenesis are manifest in an imprint that can persist in both somatic and germline gene expression for multiple generations.     

Optimal seasonal schedules and the relative dominance of heteromorphic and isomorphic life cycles in macroalgae

Marine macroalgae (seaweed) show diverse life cycles. Species with a heteromorphic life cycle have a large multicellular algal body in one generation but have a very small body in the second generation of the same year. In contrast, the diploid and haploid life forms of isomorphic species have similar morphology, and these species often have more than two generations in a year. Here, we first study the optimal life cycle schedule of marine macroalgae when daily mortality changes seasonally, and then we discuss the conditions for coexistence and relative dominance of different life cycles. According to the optimal life cycle schedule, heteromorphic species tend to have a generation with a large algal body when mortality is low, and a microscopic-sized generation when mortality is high. In contrast, isomorphic species tend to mature when body size reaches a threshold value that is the same for different generations. We then examine the coexistence of the two life cycles when growth rate decreases with biomass. The model predicts that (1) at high latitudes (i.e., in strongly seasonal environments), heteromorphic species are likely to dominate over isomorphic species, and (2) species with a heteromorphic life cycle should dominate in the supratidal and upper intertidal zones where macroalgae tend to suffer high mortality, and also in the subtidal zone, where mortality is low, whereas isomorphic species are likely to be more successful when mortality is intermediate. These predictions are consistent with the observed distribution patterns of the two life cycles in macroalgae.