The effects of running activity on the reproductive axes of rodents

Access to a running wheel causes gonadal recrudescence in Syrian hamsters whose reproductive axes have been suppressed by housing them under short day lengths (Borer et al. 1983). The first experiment tested the generality of this phenomenon in a population of rodents that is genetically heterogeneous for reproductive photoresponsiveness. Male meadow voles (Microtus pennsylvanicus) of the two extreme phenotypes — reproductively photoresponsive and non-responsive — were either provided with a running wheel or housed without one. After 4 weeks with a wheel, the responsive voles had recovered full reproductive function, while the reproductive axes of responsive voles housed without wheels remained suppressed. Three experiments queried whether the use of a wheel would have reproductively stimulative effects in other rodents. First, intact male mice given access to wheels showed no increase in testis size when compared to mice housed without wheels. Likewise, locomotor activity had no effect on male rats whose testes were partially regressed in response to testosterone implants or on female mice whose estrous cycles were pheromonally suppressed by housing them in groups. Thus the neuroendocrine pathway used by locomotor activity to enhance the secretion of gonadotropin is specifically allied with the pathway used by photoperiod to control GnRH secretion.Abbreviations GnRH gonadotropin-releasing hormone - LH luteinizing hormone     

Tau differences between short-day responsive and short-day nonresponsive white-footed mice (Peromyscus leucopus) do not affect reproductive photoresponsiveness

In laboratory-bred rodent populations, intraspecific variation in circadian system organization is a known cause of individual variation in reproductive photoresponsiveness. The authors sought to determine whether circadian system variation accounted for individual variation in reproductive photoresponsiveness in a single, highly genetically variable population of Peromyscus leucopus recently derived from the wild. Running-wheel activity patterns of male and female mice, aged 70 to 90 days, from artificially selected lines of reproductively photoresponsive (R) and nonresponsive (NR) lines were monitored under short-day photoperiod (8 h light, 16 h dark), long-day photoperiod (16 h light, 8 h dark), and constant darkness (DD). NR mice displayed a significantly longer mean free-running period (24.08 h) in DD compared with R mice (23.75 h), due in large part to a difference between NR and R females (24.25 h vs. 23.74 h, respectively). All other entrainment characteristics (alpha, phase angle of activity) under short days, long days, and DD were similar between R and NR mice. Variation in free-running period and entrainment characteristics has been shown to affect photoresponsiveness in other rodent species by altering the manner in which the circadian system interprets short days. To determine whether variation in photoresponsiveness in P. leucopus is due to differences in free-running period instead of variation downstream from the central circadian clock in the pathway controlling photoresponsiveness, the authors exposed young R and NR mice to DD and measured the effect on reproductive organ development. If variation in free-running period affected how the circadian system of mice interpreted short days, then both R and NR mice exposed to DD should have exhibited a delay in gonadal development. Only R mice exhibited pubertal delay in DD. NR mice exhibited large paired testes, paired seminal vesicles, paired ovaries, and uterine weight typical of mice nonresponsive to short days, whereas R mice exhibited reproductive organ weight typical of mice responsive to short days. These data suggest that despite significant differences in free-running period between R and NR mice, individual variation in photoresponsiveness is not due to differences in how the circadian systems of R and NR mice interpret the LD cycle.     

Evidence for genetic variation in the occurrence of the photoresponse of the Djungarian hamster, Phodopus sungorus.

The Djungarian hamster generally responds to a short-day photoperiod with a complex syndrome of physiological and behavioral changes; however, not all hamsters are photoresponsive. The phenotypic difference is, in part, genetically determined. Parent-offspring regression on a number of continuous and discontinuous measures indicated significant heritability for photoresponsiveness. Four generations of replicated bidirectional selection on a photoresponse index (PI) resulted in significant shifts in the percentage of responsive hamsters, although the average PI of responsive individuals was not significantly changed. Eight estimates of heritability ranged from 0.20 to 0.52. We hypothesize that the circadian system is responsible for the occurrence of the photoresponse, but that the extent of photoresponse is controlled by a separate functional system.     

Lack of reproductive photoresponsiveness and correlative failure to respond to melatonin in a tropical rodent, the cane mouse.

Cane mice (Zygodontomys brevicauda) are year-round breeders in Venezuela. As shown previously, these animals are not reproductively responsive to variation in photoperiod. In the present experiments, male cane mice were maintained on long or short day lengths (16L:8D or 8L:16D, respectively) and challenged with each of three experimental treatments known to "unmask" reproductive photoresponsiveness in laboratory rats: olfactory bulbectomy, prolonged food restriction, and exposure as neonates to a single injection of testosterone. Variation in photoperiod had no inhibitory effect on the responses of cane mice to any of these three treatments, as assessed by the weight of their testes and seminal vesicles. A fourth experiment demonstrated that cane mice are insensitive to 10 wk of continuous exposure to pharmacological levels of melatonin, again as assessed by reproductive organ weight. Likewise, a fifth experiment documented a lack of response to 10 wk of late-afternoon injections of massive amounts of melatonin. The cane mouse apparently is unique among the animals challenged so far in these ways in that it seems to have no vestige of reproductive photoresponsiveness.     

Response to selection for photoperiod responsiveness on the density and location of mature GnRH-releasing neurons

Natural variation in neuroendocrine traits is poorly understood, despite the importance of variation in brain function and evolution. Most rodents in the temperate zones inhibit reproduction and other nonessential functions in short winter photoperiods, but some have little or no reproductive response. We tested whether genetic variability in reproductive seasonality is related to individual differences in the neuronal function of the gonadotropin-releasing hormone network, as assessed by the number and location of mature gonadotropin-releasing hormone-secreting neurons under inhibitory and excitatory photoperiods. The experiments used lines of Peromyscus leucopus previously developed by selection from a wild population. One line contained individuals reproductively inhibited by short photoperiod, and the other line contained individuals nonresponsive to short photoperiod. Expression of mature gonadotropin-releasing hormone (GnRH) immunoreactivity in the brain was detected using SMI-41 antibody in the single-labeled avidin-biotin-peroxidase-complex method. Nonresponsive mice had 50% more immunoreactive GnRH neurons than reproductively inhibited mice in both short- and long-day photoperiods. The greatest differences were in the anterior hypothalamus and preoptic areas. In contrast, we detected no significant within-lines differences in the number or location of immunoreactive GnRH neurons between photoperiod treatments. Our data indicate that high levels of genetic variation in a single wild population for a specific neuronal trait are related to phenotypic variation in a life history trait, i.e., winter reproduction. Variation in GnRH neuronal activity may underlie some of the natural reproductive and life history variation observed in wild populations of P. leucopus. Similar genetic variation in neuronal traits may be present in humans and other species.     

Differential reproductive response to short photoperiod in deer mice: role of melatonin

Summary Inhibitory photoperiod differentially effects reproduction in deer mice (Peromyscus maniculatus nebrascensis). Pituitary-testicular function is arrested in about one-third of short-day exposed males (reproductively responsive mice), while an equal number remain fertile (reproductively nonresponsive mice). Both phenotypes are found in natural populations and their disparate reproductive responses have a genetic basis. To assess whether this difference is attributable to a prepineal/pineal or post-pineal mechanism, we compared spermatogenic responses of known and unknown phenotype to exogenous melatonin. Melatonin significantly reduced mean sperm number in long-day housed mice of unknown phenotype. But, individual responses ranged from azoospermia to normal spermatogenesis, and this range was not significantly different from that previously recorded for short-day exposed mice. Reproductively nonresponsive males were unaffected by melatonin administration when housed under long or short daylength. In contrast, melatonin significantly suppressed sperm production in reproductively responsive males housed under long photoperiod, but had no additional suppressive effect in short-day housed mice with regressed testes. These data demonstrate that melatonin is only effective in eliciting testicular regression in reproductively responsive males. Taken together, these results suggest that differential testicular response to photoperiod are caused by a post-pineal mechanism.Abbreviations LD long day - SD short day - 16L:8D 16 h light, 8 h dark - 8L:16D 8 h light, 16 h dark     

Photoperiod, melatonin secretion, and sexual maturation in a tropical rodent

Descendants of a sample of cane mice (Zygodontomys brevicauda) trapped at 8 degrees latitude in Venezuela were tested for reproductive photoresponsiveness. This species breeds continuously, year around, despite living in a seasonally harsh habitat. At 50 days of age there were no differences in the weights of the testes or seminal vesicles or in sperm counts of males born and reared on 16L:8D, 13L:11D, 11L:13D, or 8L:16D photoperiods, although there were small differences in body weight. Females born and reared on 16L:8D vs. 8L:16D cycles became pregnant at the same rates and ages when paired with males at 21 or 31 days of age. The daily duration of melatonin secretion depended on the length of the dark phase of the cycle in both sexes. Circulating levels of melatonin were elevated for 8 h on a 16L:8D cycle and for between 9 and 16 h on an 8L:16D cycle. In this tropical species, the neuroendocrine pathway that links photoperiod to reproduction apparently is disconnected somewhere between melatonin and gonadotropin secretion, causing cane mice to be reproductively unresponsive to variation in photoperiod.     

Are humans seasonally photoperiodic?

Humans exhibit seasonal variation in a wide variety of behavioral and physiological processes, and numerous investigators have suggested that this might be because we are sensitive to seasonal variation in day length. The evidence supporting this hypothesis is inconsistent. A new hypothesis is offered here-namely, that some humans indeed are seasonally photoresponsive, but others are not, and that individual variation may be the cause of the inconsistencies that have plagued the study of responsiveness to photoperiod in the past. This hypothesis is examined in relation to seasonal changes in the reproductive activity of humans, and it is developed by reviewing and combining five bodies of knowledge: correlations of human birthrates with photoperiod; seasonal changes in the activity of the neuroendocrine pathway that could link photoperiod to gonadal steroid secretion in humans; what is known about photoperiod, latitude, and reproduction of nonhuman primates; documentation of individual variation in photoresponsiveness in rodents and humans; and what is known about the evolutionary ecology of humans.     

Photoperiodic polyphenisms in rodents: neuroendocrine mechanisms, costs, and functions

Annual changes in daylength figure prominently in the generation of seasonal rhythms in reproduction, and a wide variety of mammals use ambient photoperiod as a proximate cue to time critical reproductive events. Nevertheless, within many reproductively photoperiodic mammalian species, there exist individuals--termed "photoperiod nonresponders"--that fail to adopt a seasonal breeding strategy and instead exhibit reproductive competence at a time of year when their conspecifics are reproductively quiescent. Photoperiod nonresponsiveness has been principally characterized by laboratory observations--over half of the species known to be reproductively photoperiodic contain a proportion of nonresponsive individuals. The study of nonresponders has generated basic insights regarding photic regulation of reproduction in mammals. The neuroendocrine mechanisms by which the short-day photoperiodic signal is degraded or lost in nonresponders varies between species: differences in features of the circadian pacemaker, which provides photoperiodic input to the reproductive neuroendocrine system, have been identified in hamsters; changes in the responsiveness of hypothalamic gonadotrophs to melatonin and as-yet-unspecified inhibitory signals have been implicated in voles and mice. Individuals that continue to breed when their conspecifics refrain might enjoy higher fitness under certain circumstances. Statements regarding the adaptive function of reproductive nonresponsiveness to photoperiod require additional information on the costs (metabolic and fitness) of sustaining reproductive function during the winter months and how these costs vary as a function of environmental conditions. Reproductive nonresponders thus continue to represent a challenge to theories that extol the adaptive function of seasonality. Several nonexclusive hypotheses are proposed to account for the maintenance of nonresponsive individuals in wild rodent populations.     

Phenotypic differences in the GnRH neuronal system of deer mice Peromyscus maniculatus under a natural short photoperiod

The neural mechanisms by which short photoperiod induces gonadal regression among seasonally breeding mammals are not well understood. One hypothesis suggests that the proximate cause of seasonal gonadal regression is a photoperiod-induced modification in GnRH secretion. This hypothesis is indirectly supported by our recent findings using immunocytochemistry which identified specific photoperiod-induced adjustments in the number and morphology of GnRH containing neurones between reproductively competent and reproductively regressed laboratory housed male deer mice. Herein, we report that the GnRH neuronal system is similarly affected in reproductively responsive and nonresponsive wild male deer mice Peromyscus maniculatus exposed to a natural short photoperiod. The distribution of immunoreactive (IR)-GnRH neurones was nearly identical in field caught animals and those housed under artificial photoperiod in the laboratory. Compared with reproductively nonresponsive males, reproductively responsive mice from the field population possessed a greater total number of IR-GnRH neurones, a greater number of IR-GnRH neurones within the lateral hypothalamus, and a greater proportion of bipolar IR-GnRH neurones. Each of these distributional and morphological characters was consistent with our findings in laboratory housed male deer mice exposed to an artificial short photoperiod. Taken together, these data underscore the validity of using an artificial photoperiod to evaluate seasonal adjustments in reproductive function in the laboratory.     

Genetic analyses of photoresponsiveness in the Djungarian hamster,Phodopus sungorus

Summary Endotherms living at temperate and arctic latitudes must adjust their physiology and behavior in order to survive seasonal change. The Djungarian hamster uses photoperiod to cue annual cycles of reproduction and thermoregulation, and its responses to short photoperiod include loss of body weight and change in pelage color. Some individuals do not exhibit these responses when exposed to short days.In this study individual variation in photoresponsiveness is quantified, and four lines of evidence for a genetic component to that variation are provided. First, two separate breeding stocks differed in both the percent of animals responding to a short-day lighting regimen (SD) and in the degree and timing of their response. Second, analysis of variance within and between families of full sibs for a photoresponsive index, PI (body weightloss +2 (molt index –1)) following 12 weeks in SD demonstrated a significant family resemblance (intraclass correlation of 0.36±0.03). Third, heritability estimates from regression of offspring scores on parent scores for body weight loss, molt index and PI after 12 weeks in SD were 0.340.13, 0.36±0.10 and 0.37±0.12, respectively, indicating a strong additive genetic component for the three characters. Finally, a significant response occurred after one generation of artificial selection for and against photoresponsiveness.Abbreviations PI photoresponsive index (body weight loss+2 (molt index –1)) - SD short day light regimen     

Patterns of parapatric speciation

Abstract. Geographic variation may ultimately lead to the splitting of a subdivided population into reproductively isolated units in spite of migration. Here, we consider how the waiting time until the first split and its location depend on different evolutionary factors including mutation, migration, random genetic drift, genetic architecture, and the geometric structure of the habitat. We perform large-scale, individual-based simulations using a simple model of reproductive isolation based on a classical view that reproductive isolation evolves as a by-product of genetic divergence. We show that rapid parapatric speciation on the time scale of a few hundred to a few thousand generations is plausible even when neighboring subpopulations exchange several individuals each generation. Divergent selection for local adaptation is not required for rapid speciation. Our results substantiates the claims that species with smaller range sizes (which are characterized by smaller local densities and reduced dispersal ability) should have higher speciation rates. If mutation rate is small, local abundances are low, or substantial genetic changes are required for reproductive isolation, then central populations should be the place where most splits take place. With high mutation rates, high local densities, or with moderate genetic changes sufficient for reproductive isolation, speciation events are expected to involve mainly peripheral populations.     

Time scales of divergence and speciation among natural populations and subspecies of Arabidopsis lyrata (Brassicaceae)

? Premise of the study: Plant populations that face new environments adapt and diverge simultaneously, and both processes leave footprints in their genetic diversity. Arabidopsis lyrata is an excellent species for studying these processes. Pairs of populations and subspecies of A. lyrata represent different stages of divergence. These populations are also known to be locally adapted and display various stages of emerging reproductive isolation. ? Methods: We used nucleotide diversity data from 19 loci to estimate divergence times and levels of diversity among nine A. lyrata populations. Traditional distance-based methods and model-based clustering analysis were used to supplement pairwise coalescence-based analysis of divergence. ? Key results: Estimated divergence times varied from 130000 generations between North American and European subspecies to 39000 generations between central European and Scandinavian populations. In concordance with previous studies, the highest level of diversity was found in Central Europe and the lowest in North America and a diverged Russian Karhum?ki population. Local adaptation among Northern and central European populations has emerged during the last 39000 generations. Populations that are reproductively isolated by prezygotic mechanisms have been separated for a longer time period of ～70000 generations but still have shared nucleotide polymorphism. ? Conclusions: In A. lyrata, reproductively isolated populations started to diverge ～70000 generations ago and more closely related, locally adapted populations have been separate lineages for ～39000 generations. However, based on the posterior distribution of divergence times, the processes leading to reproductive isolation and local adaptation are likely to temporally coincide.     

Seasonal change in the opportunity for sexual selection

Environmental and population parameters that influence the strength of sexual selection may vary considerably over the course of the reproductive season. However, the potential for sexual selection frequently fails to translate into variation in reproductive success among individuals. We investigated seasonal changes in variation in reproductive success, measured as the opportunity for sexual selection, using parentage analysis in 20 experimental populations of the European bitterling (Rhodeus amarus, Cyprinidae), a small freshwater fish with a promiscuous, resource-based mating system. We showed that although the largest males sired most offspring over the entire reproductive season, variation in reproductive success and hence the opportunity for sexual selection was low at the start of the season but increased significantly at its end. This seasonal difference probably arose from the superior competitive endurance of large males and from a higher temporal clustering of reproductively active females at the start of the breeding season than later in the season. The spatial distribution of oviposition sites had a negligible effect on the variation in reproductive success. We discuss the potential implications of our results for the importance and strength of sexual selection in natural populations.     

Genetic variation for seasonal adaptation in Peromyscus leucopus: nonreciprocal breakdown in a population cross

The genetic basis for adaptations to the diverse environments encountered by a wide-ranging species should be reflected in the phenotypes of hybrids between differentiated populations. We crossed mice from two ecologically different populations to determine whether adaptations to seasonality in Peromyscus leucopus, the white-footed mouse, display directional dominance or whether they are affected by specific interactions between genes. Connecticut mice (C) have many adaptations to seasonality that are reduced or absent in Georgia mice (G) and that affect both reproductive and thermoregulatory traits. Because these adaptations are cued by photoperiod, temperature, or both, parental and hybrid mice were acclimated to 13 degrees C in either long-day or short-day photoperiod, and several morphological, reproductive, and thermoregulatory traits were measured. Several traits, especially those involved in response to the environment, such as nest size at 13 degrees C, reproductive regression under short-day photoperiod, and molt to winter pelage, showed a non-Mendelian pattern of inheritance. However, there was a pronounced difference between maternal lineages in the F2 generation in that C-maternal F2 mice were less responsive to short-day photoperiod and cold than were the G-maternal F2 mice. Because these two classes of F2 mice are genetically equivalent, this breakdown cannot be explained as a disruption of epistatic gene interactions unless recombination rates are higher in the C-maternal lineage.     

Hypothyroidism in reproductively inhibited prairie deer mice (Peromyscus maniculatus bairdi) from laboratory populations

Several thyroid function parameters were compared between reproductively inhibited prairie deer mice of both sexes taken from laboratory populations and corresponding reproductively capable controls. The results of these experiments indicated the following: 1) prairie deer mice females had a statistically significant daily variation in mean serum thyroxine concentration and males displayed a similar trend; 2) total serum thyroxine and triiodothyronine were significantly lower in both male and female reproductively inhibited population animals compared with reproductively capable controls; 3) several morphometric characteristics of the thyroid of male and female population prairie deer mice were significantly different from that of control males and females, suggesting functional hypothyroidism in both sexes; 4) thyroid histology of male population deer mice was different from that of female population animals. In males, the data suggested that thyroid-stimulating hormone (TSH) stimulation was deficient due to some undetermined secondary hypothyroidism. In females, reduced serum thyroxine and triiodothyronine concentrations were observed due to primary hypothyroidism of unknown origin. The interrelationship between thyroid and adrenal function is discussed here and it is concluded that each of the systems may contribute to the observed reproductive inhibition. In particular, it was noted that the mechanism of response between population males and females may be quite different. No conclusive data are available to suggest whether one or the other system is the primary controller of the reproductive inhibition.     

The Evolution and Consequences of Sex-Specific Reproductive Variance

Natural selection favors alleles that increase the number of offspring produced by their carriers. But in a world that is inherently uncertain within generations, selection also favors alleles that reduce the variance in the number of offspring produced. If previous studies have established this principle, they have largely ignored fundamental aspects of sexual reproduction and therefore how selection on sex-specific reproductive variance operates. To study the evolution and consequences of sex-specific reproductive variance, we present a population-genetic model of phenotypic evolution in a dioecious population that incorporates previously neglected components of reproductive variance. First, we derive the probability of fixation for mutations that affect male and/or female reproductive phenotypes under sex-specific selection. We find that even in the simplest scenarios, the direction of selection is altered when reproductive variance is taken into account. In particular, previously unaccounted for covariances between the reproductive outputs of different individuals are expected to play a significant role in determining the direction of selection. Then, the probability of fixation is used to develop a stochastic model of joint male and female phenotypic evolution. We find that sex-specific reproductive variance can be responsible for changes in the course of long-term evolution. Finally, the model is applied to an example of parental-care evolution. Overall, our model allows for the evolutionary analysis of social traits in finite and dioecious populations, where interactions can occur within and between sexes under a realistic scenario of reproduction.     

Mechanisms for delayed density-dependent reproductive traits in field voles, Microtus agrestis: the importance of inherited environmental effects

Reproductive traits of voles vary with the phases of the population density fluctuations. We sought to determine whether the source of this variation resides in the individuals or in their environment. Overwintering field voles ( Microtus agrestis ) from two cyclic out-of-phase populations (increase and peak phases) were sampled in early spring and bred in the laboratory for two generations under standardised conditions with ambient light and temperature. Monitoring of the source populations by capture-mark-recapture showed large differences in reproductive performance. In the increase area, reproduction started six weeks earlier, the probability of maturation of young-of-the-year was more than ten times higher during mid-summer, and reproduction continued nearly two months later in the autumn than in the peak area. These differences were not found to be associated with a difference in age structure of overwintered animals between the two areas (assessed by the distribution of eye lens masses from autopsy samples). Although the population differences in reproductive traits were to some degree also present among the overwintered animals in the laboratory, we found no difference in reproductive traits in the laboratory-born generations. There was a strongly declining seasonal trend in probability of sexual maturation both in the field and in the laboratory under ambient light conditions. However, in the field there were large population differences in the steepness of the seasonal decline that were not seen under the standardised laboratory conditions. We conclude that seasonal decline in maturation rates is governed by change in photoperiod, but that the population level variation in the shape of the decline is caused by a direct response to the environment and not due to variation in any intrinsic state of the individuals reflecting the environment experienced by the previous generation(s).     

DNA barcoding will often fail to discover new animal species over broad parameter space

With increasing force, genetic divergence of mitochondrial DNA (mtDNA) is being argued as the primary tool for discovery of animal species. Two thresholds of single-gene divergence have been proposed: reciprocal monophyly, and 10 times greater genetic divergence between than within species (the "10x rule"). To explore quantitatively the utility of each approach, we couple neutral coalescent theory and the classical Bateson-Dobzhansky-Muller (BDM) model of speciation. The joint stochastic dynamics of these two processes demonstrate that both thresholds fail to "discover" many reproductively isolated lineages under a single incompatibility BDM model, especially when BDM loci have been subject to divergent selection. Only when populations have been isolated for > 4 million generations did these thresholds achieve error rates of < 10% under our model that incorporates variable population sizes. The high error rate evident in simulations is corroborated with six empirical data sets. These properties suggest that single-gene, high-throughput approaches to discovering new animal species will bias large-scale biodiversity surveys, particularly toward missing reproductively isolated lineages that have emerged by divergent selection or other mechanisms that accelerate reproductive isolation. Because single-gene thresholds for species discovery can result in substantial error at recent divergence times, they will misrepresent the correspondence between recently isolated populations and reproductively isolated lineages (= species).     

Partial interfertility between independently originated populations of the neo-allopolyploid <Emphasis Type="Italic">Mimulus peregrinus</Emphasis>

The reduction in genetic diversity in polyploid lineages, formed from whole-genome duplication of a few individuals, can affect their long-term evolutionary potential. Because most polyploids originate multiple times, secondary contact and gene exchange among independently formed polyploids can create novel genetic combinations and reduce the severity of genetic bottlenecks. However, independently originated polyploids may be reproductively isolated from each other due to genetic and chromosomal differences predating the polyploidisation event, or evolving subsequently in the dynamic genomes of young polyploid populations. Here we conducted experimental crosses to investigate the phenotype and interfertility between two independently originated populations of the allopolyploid Mimulus peregrinus (Phrymaceae). We found that individuals from the two populations are phenotypically distinct, but that inter- and intrapopulation crosses are not statistically different. Interpopulation crosses produce viable and fertile offspring, although our results suggest the existence of partial reproductive barriers in the form of reduced pollen viability. We found no difference in pollen viability between F1 and F2 generations. In contrast, we detected a reduction in floral and vegetative size, and in the proportion of plants that flowered, between F1 and F2 generations for both intra- and interpopulation crosses. Together, our results indicate that populations of independent origin can partially exchange genes, producing variable offspring, and that the phenotype of M. peregrinus may be unstable in the early generations. Natural selection on genetically based variation may be required for the evolution of more stable and fertile individuals of this nascent allopolyploid.