Comparative genetics of potential prezygotic and postzygotic isolating barriers in a Lycopersicon species cross

I compare the genetic basis of quantitative traits that potentially contribute to pre- and postzygotic isolation between the plant species Solanum lycopersicum (formerly Lycopersicon esculentum) and Solanum habrochaites (formerly Lycopersicon hirsutum), using quantitative trait loci (QTL) mapping in a set of near-isogenic lines. Putative prezygotic isolating traits include flower size, flower shape, stigma exertion, and inflorescence length, that can influence pollinator preferences and/or selfing rates, and therefore gene flow between divergent types. Postzygotic isolating traits are hybrid pollen and seed sterility. Three substantive results emerge from these analyses. First, the genetic basis of floral differentiation appears to be somewhat less complex than the genetic basis of postzygotic hybrid sterility, although these differences are very modest. Second, there is little evidence that traits for floral differentiation are causally or mechanistically associated with hybrid sterility traits in this species cross. Third, there is little evidence that hybrid sterility QTL are more frequently associated with chromosomal centromeric regions, in comparison to floral trait QTL, a prediction of centromeric drive models of hybrid sterility. Although genome-wide associations are not evident in this analysis, several individual chromosomal regions that contain clusters of QTL for both floral and sterility traits, or that indicate hybrid sterility effects at centromere locations, warrant further fine-scale investigation.     

Strong asymmetry in the relative strengths of prezygotic and postzygotic barriers between two damselfly sister species

One of the longest debates in biology has been over the relative importance of different isolating barriers in speciation. However, for most species, there are few data evaluating their relative contributions and we can only speculate on the general roles of pre- and postzygotic isolation. Here, we quantify the absolute and cumulative contribution of 19 potential reproductive barriers between two sympatric damselfly sister species, Ischnura elegans and I. graellsii, including both premating (habitat, temporal, sexual and mechanical isolation) and postmating barriers (prezygotic: sperm insemination success and removal rate, oviposition success, fertility, fecundity; postzygotic: hybrid viability, hybrid sterility and hybrid breakdown). In sympatry, total reproductive isolation between I. elegans females and I. graellsii males was 95.2%, owing mostly to a premating mechanical incompatibility (93.4%), whereas other barriers were of little importance. Isolation between I. graellsii females and I. elegans males was also nearly complete (95.8%), which was caused by the cumulative action of multiple prezygotic (n= 4, 75.4%) and postzygotic postmating barriers (n= 5, 7.4%). Our results suggest that premating barriers are key factors in preventing gene flow between species, and that the relative strengths of premating barriers is highly asymmetrical between the reciprocal crosses.     

Geographical variation in postzygotic isolation and its genetic basis within and between two Mimulus species

The aim of this study is to investigate the evolution of intrinsic postzygotic isolation within and between populations of Mimulus guttatus and Mimulus nasutus. We made 17 intraspecific and interspecific crosses, across a wide geographical scale. We examined the seed germination success and pollen fertility of reciprocal F₁ and F₂ hybrids and their pure-species parents, and used biometrical genetic tests to distinguish among alternative models of inheritance. Hybrid seed inviability was sporadic in both interspecific and intraspecific crosses. For several crosses, Dobzhansky–Muller incompatibilities involving nuclear genes were implicated, while two interspecific crosses revealed evidence of cytonuclear interactions. Reduced hybrid pollen fertility was found to be greatly influenced by Dobzhansky–Muller incompatibilities in five out of six intraspecific crosses and nine out of 11 interspecific crosses. Cytonuclear incompatibilities reduced hybrid fitness in only one intraspecific and one interspecific cross. This study suggests that intrinsic postzygotic isolation is common in hybrids between these Mimulus species, yet the particular hybrid incompatibilities responsible for effecting this isolation differ among the populations tested. Hence, we conclude that they evolve and spread only at the local scale.     

Multiple strong postmating and intrinsic postzygotic reproductive barriers isolate florally diverse species of Jaltomata (Solanaceae)

Divergence in phenotypic traits often contributes to premating isolation between lineages, but could also promote isolation at postmating stages. Phenotypic differences could directly result in mechanical isolation or hybrids with maladapted traits; alternatively, when alleles controlling these trait differences pleiotropically affect other components of development, differentiation could indirectly produce genetic incompatibilities in hybrids. Here, we determined the strength of nine postmating and intrinsic postzygotic reproductive barriers among 10 species of Jaltomata (Solanaceae), including species with highly divergent floral traits. To evaluate the relative importance of floral trait diversification for the strength of these postmating barriers, we assessed their relationship to floral divergence, genetic distance, geographical context, and ecological differences, using conventional tests and a new linear‐mixed modeling approach. Despite close evolutionary relationships, all species pairs showed moderate to strong isolation. Nonetheless, floral trait divergence was not a consistent predictor of the strength of isolation; instead this was best explained by genetic distance, although we found evidence for mechanical isolation in one species, and a positive relationship between floral trait divergence and fruit set isolation across species pairs. Overall, our data indicate that intrinsic postzygotic isolation is more strongly associated with genome‐wide genetic differentiation, rather than floral divergence.     

When interspecific prezygotic barriers break down: hybridization between two Potamogeton species (P. wrightii and P. perfoliatus) and comparison of the artificial and natural hybrids

Interspecific hybridization played an important role in speciation and evolution of angiosperms. Although the widespread occurrence of natural hybrids in the genus Potamogeton has been studied intensively, few successful experimental hybridization studies have been reported in this genus. In the present study, critical experimental hybridization was conducted using Potamogeton?×?intortusifolius, a natural hybrid widely distributed in China, and its parents (P. perfoliatus and P. wrightii). The absence of prezygotic barriers between P. wrightii and P. perfoliatus was observed, which contributes to the frequent hybridization of these species in nature. The pollen tube growth rates of P. perfoliatus were much faster than those of P. wrightii in the style of that species. However, the conspecific pollen tubes were competitively advantageous in P. perfoliatus styles. The interspecific hybridization could be applied bidirectionally, and 28 F1 hybrid individuals were successfully obtained from P. wrightii?×?P. perfoliatus, despite the low germination possibility of the hybrid seeds. Both the artificial and natural hybrids exhibited intermediate morphological characters but presented much lower fertility. The sterility of P.?×?intortusifolius is mainly due to its low female fitness. However, the offspring from P. wrightii?×?P.?×?intortusifolius indicated the potential for backcrossing in nature. This is the most successful attempt at artificial hybridization in this genus so far. The possible route for restoration of fertility and the fitness of the hybrids are also discussed in this paper.     

Pre‐ and postzygotic mechanisms preventing hybridization in co‐occurring species of the Impatiens purpureoviolacea complex

In the species‐rich genus Impatiens, few natural hybrids are known, even though closely related species often occur sympatrically. In this study, we aim to bridge the gap between micro‐ and macro‐evolution to disentangle pre‐ and postzygotic mechanisms that may prevent hybridization in the Impatiens purpureoviolacea complex from Central Africa. We analyzed habitat types, species distribution, pollination syndromes, pollinator dependency, genome sizes, and chromosome numbers of seven out of the ten species of the complex as well as of one natural hybrid and reconstructed the ancestral chromosome numbers of the complex. Several species of the complex occur in sympatry or geographically very close to each other. All of them are characterized by pre‐ and/or postzygotic mechanisms potentially preventing hybridization. We found four independent polyploidization events within the complex. The only known natural hybrid always appears as single individual and is self‐fertile. But the plants resulting from self‐pollinated seeds often die shortly after first flowering. These results indicate that the investigated mechanisms in combination may effectively but not absolutely prevent hybridization in Impatiens and probably occur in other genera with sympatric species as well.     

Intraspecific variation in reproductive barriers between two closely related Arabidopsis sister species

Reproductive isolation (RI) is a critical component of speciation and varies strongly in timing and strength among different sister taxa, depending on, for example the geography of speciation and divergence time. However, these factors may also produce variation in timing and strength among populations within species. Here we tested for variation in the expression of RI among replicate population pairs between the sister taxa Arabidopsis lyrata subsp. lyrata and A. arenicola. While the former is predominantly outcrossing, the latter is predominantly selfing. We focused on intrinsic prezygotic and postzygotic RI as both species occur largely in allopatry. We assessed RI by performing within-population crosses and interspecific between-population crosses, and by raising offspring. RI was generally high between all interspecific population pairs, but it varied in timing and strength depending on population history. Prezygotic isolation was strongest between the closest-related population pair, while early postzygotic isolation was high for all other population pairs. Furthermore, the timing and strength of RI depended strongly on cross direction. Our study provides empirical support that reproductive barriers between species are highly variable among population pairs and asymmetric within population pairs, and this variation seems to follow patterns typically described across species pairs.     

Unidirectional hybridization and reproductive barriers between two heterostylous primrose species in north-west Yunnan,China

Background and Aims

Heteromorphy in flowers has a profound effect on breeding patterns within a species, but little is known about how it affects reproductive barriers between species. The heterostylous genus Primula is very diverse in the Himalaya region, but hybrids there have been little researched. This study examines in detail a natural hybrid zone between P. beesiana and P. bulleyana.

Methods

Chloroplast sequencing, AFLP (amplified fragment length polymorphism) markers and morphological comparisons were employed to characterize putative hybrids in the field, using synthetic F₁s from hand pollination as controls. Pollinator visits to parent species and hybrids were observed in the field. Hand pollinations were conducted to compare pollen tube growth, seed production and seed viability for crosses involving different morphs, species and directions of crossing.

Key Results

Molecular data revealed all hybrid derivatives examined to be backcrosses of first or later generations towards P. bulleyana: all had the chloroplast DNA (cpDNA) of this species. Some individuals had morphological traits suggesting they were hybrids, but they were genetically similar to P. bulleyana; they might have been advanced generation backcrosses. Viable F₁s could not be produced with P. bulleyana pollen on P. beesiana females, irrespective of the flower morphs used. Within-morph crosses for each species had very low (<10 %) seed viability, whereas crosses between pin P. bulleyana (female) and pin P. beesiana had a higher seed viability of 30 %. Thus genetic incompatibility mechanisms back up mechanical barriers to within-morph crosses in each species, but are not the same between the two species. The two species share their main pollinators, and pollinators were observed to fly between P. bulleyana and hybrids, suggesting that pollinator behaviour may not be an important isolating factor.

Conclusions

Hybridization is strongly asymmetric, with P. bulleyana the only possible mother and all detected hybrids being backcrosses in this direction. Partial ecological isolation and inhibition of heterospecific pollen, and possibly complete barriers to F₁ formation on P. beesiana, may be enough to make F₁ formation very rare in these species. Therefore, with no F₁ detected, this hybrid zone may have a finite life span as successive generations become more similar to P. bulleyana.     

Hybrid asexuality as a primary postzygotic barrier between nascent species: On the interconnection between asexuality,hybridization and speciation

Although sexual reproduction is ubiquitous throughout nature, the molecular machinery behind it has been repeatedly disrupted during evolution, leading to the emergence of asexual lineages in all eukaryotic phyla. Despite intensive research, little is known about what causes the switch from sexual reproduction to asexuality. Interspecific hybridization is one of the candidate explanations, but the reasons for the apparent association between hybridization and asexuality remain unclear. In this study, we combined cross‐breeding experiments with population genetic and phylogenomic approaches to reveal the history of speciation and asexuality evolution in European spined loaches (Cobitis). Contemporary species readily hybridize in hybrid zones, but produce infertile males and fertile but clonally reproducing females that cannot mediate introgressions. However, our analysis of exome data indicates that intensive gene flow between species has occurred in the past. Crossings among species with various genetic distances showed that, while distantly related species produced asexual females and sterile males, closely related species produce sexually reproducing hybrids of both sexes. Our results suggest that hybridization leads to sexual hybrids at the initial stages of speciation, but as the species diverge further, the gradual accumulation of reproductive incompatibilities between species could distort their gametogenesis towards asexuality. Interestingly, comparative analysis of published data revealed that hybrid asexuality generally evolves at lower genetic divergences than hybrid sterility or inviability. Given that hybrid asexuality effectively restricts gene flow, it may establish a primary reproductive barrier earlier during diversification than other “classical” forms of postzygotic incompatibilities. Hybrid asexuality may thus indirectly contribute to the speciation process.     

DNA hybridization to compare species compositions of natural bacterioplankton assemblages

Little is known about the species composition and variability of natural bacterial communities, mostly because conventional identification requires pure cultures, but less than 1% of active natural bacteria are cultivable. This problem was circumvented by comparing species compositions via hybridization of total DNA of natural bacterioplankton communities for the estimation of the fraction of DNA in common between two samples (similarity). DNA probes that were labeled with 35S by nick translation were hybridized to filter-bound DNA in a reciprocal fashion; similarities (in percent) were calculated by normalizing the values to self-hybridizations. In tests with DNA mixtures of pure cultures, the experimentally observed similarities agreed with expectations. However, reciprocal similarities (probe and target reversed) were often asymmetric, unlike those of DNA from single strains. This was due to the relative complexity and G + C content of DNA, which provided a means to interpret the asymmetry that was occasionally observed in natural samples. Natural bacteria were collected by filtration from Long Island Sound (LIS), N.Y., the Caribbean and Sargasso seas, and a coral reef lagoon near Bermuda. The samples showed similarities of less than 10 to 95%. The LIS and Sargasso and Caribbean sea samples were 20 to 50% similar to each other. The coral reef sample was less than 10% similar to the others, indicating its unique composition. Seasonality was also observed; an LIS sample obtained in the autumn was 40% similar to two LIS samples obtained in the summer; these latter two samples were 95% similar. We concluded that total DNA hybridization is a rapid, simple, and unbiased method for investigating the variation of bacterioplankton species composition over time and space, avoiding the need of culturing.     

The effects of artificial and natural barriers on the movement of small mammals in Banff National Park, Canada

Barriers that dissect continuous habitat, such as roads, initiate the process of habitat fragmentation and can filter or eliminate animal movement through otherwise pristine areas. A severe road barrier appears to exist in Banff National Park where the Trans‐Canada Highway (TCH) is known to impede the movement of some animals, but the effects of this barrier relative to natural barriers and continuous habitat have not been assessed. To compare animal movement in response to these three contexts, we translocated three species of murid rodents (meadow voles, Microtus pennsylvanicus, deer mice, Peromyscus maniculatus, and red‐backed voles, Clethrionomys gapperi) across the TCH (an artificial barrier), a forested strip in the median of the TCH (a natural barrier), in the adjacent highway verge (continuous grassy) and in nearby forest (continuous forested). We coated murids with fluorescent dye, released them at standardized distances, and followed the resulting trails to obtain detailed information on movement paths. Overall, individuals were 20% less successful crossing the TCH than natural (forested) barriers and 10% less successful crossing natural barriers than continuous habitat, but exhibited marked variation among species. Path tortuosity was negatively related to return success; more convoluted paths were associated with lower success across all treatments. Deer mice returned to their home‐ranges more often than the other two species, perhaps because of their generalist habitat preferences, nocturnal activities, or larger home ranges. In all treatments, the fractal dimensions of movement paths were very qualitatively similar to those observed for return success and tortuosity, indicating that the responses of these species were not influenced by spatial scale. Together, our results suggest that (a) murid species differ in the movement limitation imposed by both natural and artificial barriers, perhaps as a consequence of differences in habitat preferences and activity patterns, and (b) murid responses to a road barrier are only moderately different from responses to the natural barrier of a forest edge in the context of translocation experiments.     

探讨种间传粉在杜鹃花属自然杂交物种形成中的作用

自然杂交是物种形成的一个途径, 在植物进化中起着重要的作用。自然杂交主要通过种间的基因交流, 花粉传递则是基因交流的主要途径。马缨花(Rhododendron delavayi)、大白花杜鹃(R. decorum)、迷人杜鹃(R. agastum)和露珠杜鹃(R. irroratum)是在云南广泛分布的杜鹃花种类, 马缨花与大白花杜鹃形态上区别明显, 而它们的可能杂交种迷人杜鹃和露珠杜鹃在形态上基本介于二者之间。本文对这4种杜鹃花的开花物候和访花昆虫的种类进行了观察,并进行了其繁育系统和种子萌发的实验。我们发现马缨花的花期从3月初至5月底, 迷人杜鹃与露珠杜鹃花期基本一致, 为3月初至4月初, 二者同大白花杜鹃基本不存在花期重叠, 大白花杜鹃的花期为4月中旬至5月底。4种杜鹃花的开花期不同年份稍有变化, 其单花开花周期都在一周以上。4种杜鹃花的传粉昆虫虽然种类和数量上有所不同, 但都以膜翅目和双翅目昆虫为主, 中华蜜蜂(Apis cerana cerana)是其共有的传粉昆虫。繁育系统研究发现, 除露珠杜鹃外, 其余3种自花不育, 而种间杂交不存在任何生殖障碍, 可以产生萌发率很高的种子。通过分析4种杜鹃花开花物候重叠、共有传粉昆虫及种间杂交可育等现象, 探讨了传粉昆虫和异花授粉的机制在自然杂交物种形成中的作用及杂交种的适应能力。     

Introgressive hybridization as the breakdown of postzygotic isolation: a theoretical perspective

In this paper, both the empirical and theoretical genetic aspects of human-mediated introgressive hybridization are reviewed in terms of their association with the breakdown of postzygotic isolating mechanisms. I also compare several simulation models with an ecological or genetic focus that are relevant to the prediction and risk assessment of genetic extinction due to hybridization. One barrier to devising comprehensive risk assessment frameworks is a lack of sufficient population genetic studies that associate introgressive hybridization with specific isolating mechanisms. A gametic model based on multilocus underdominant fitness is one of the best genetic models for introgressive hybridization because it explicitly incorporates the postzygotic isolating mechanism known as Dobzhansky–Muller genetic incompatibility.     

天然杂交与遗传渐渗对植物入侵性的影响

生物入侵给全球生态环境与社会经济都带来了严重危害, 对其入侵机制的研究非常重要。生物入侵是一个适应性进化的过程, 天然杂交与遗传渐渗可以改变外来物种对环境的适应性并提高其入侵能力, 使其进化成为入侵种。因此了解杂交-渐渗在促进生物入侵过程中的遗传作用, 将有助于我们采取有效措施来控制生物入侵及其危害。本文从杂交-渐渗对生物适应性进化和物种形成影响的角度, 阐明外来种如何通过杂交-渐渗在新的生境中改变其适应性、生存竞争能力和入侵能力。杂交-渐渗可以导致物种发生多倍体水平和同倍体水平的进化, 虽然二者的进化过程不尽相同, 但均能使杂种群体在遗传上产生较大变化, 进而影响杂种群体的适合度, 这一过程可能促使外来种在新的生境中的成功入侵进而转变为入侵种。随着转基因生物技术的迅速发展, 大量转基因作物进入环境释放和商品化种植, 具有特定功能的转基因可能通过杂交-渐渗进入野生近缘种群体, 也可能使之成为入侵性强的农田杂草, 带来难以预测的生态后果。总之, 生物入侵是一个复杂的进化和生态过程, 利用杂交-渐渗的理论来解释植物的入侵性, 仅从一个方面反映了入侵生物学的研究, 杂交-渐渗与其他理论的结合, 将从更深的层次来解释外来种的入侵机制。     

Insights into the ecology of Schizosaccharomyces species in natural and artificial habitats

The fission yeast genus Schizosaccharomyces contains important model organisms for biological research. In particular, S. pombe is a widely used model eukaryote. So far little is known about the natural and artificial habitats of species in this genus. Finding out where S. pombe and other fission yeast species occur and how they live in their habitats can promote better understanding of their biology. Here we investigate in which substrates S. pombe, S. octosporus, S. osmophilus and S. japonicus are present. To this end about 2100 samples consisting of soil, tree sap fluxes, fresh fruit, dried fruit, honey, cacao beans, molasses and other substrates were analyzed. Effective isolation methods that allow efficient isolation of the above mentioned species were developed. Based on the frequency of isolating different fission yeast species in various substrates and on extensive literature survey, conclusions are drawn on their ecology. The results suggest that the primary habitat of S. pombe and S. octosporus is honeybee honey. Both species were also frequently detected on certain dried fruit like raisins, mango or pineapple to which they could be brought by the honey bees during ripening or during drying. While S. pombe was regularly isolated from grape mash and from fermented raw cacao beans S. octosporus was never isolated from fresh fruit. The main habitat of S. osmophilus seems to be solitary bee beebread. It was rarely isolated from raisins. S. japonicus was mainly found in forest substrates although it occurs on fruit and in fruit fermentations, too.

     

The genetics of floral divergence and postzygotic barriers between outcrossing and selfing populations of Arenaria uniflora (Caryophyllaceae)

The genetic architecture of floral traits involved in the evolution of self-pollination provides a window into past processes of mating system divergence. In this study, we use two generations of crosses between highly selfing and predominantly outcrossing populations of Arenaria uniflora (Caryophyllaceae) to determine the minimum number, average dominance relationships, and pleiotropic effects of genetic factors involved in floral divergence. Comparison of the F1 and F2 phenotypic means with the expectations of a completely additive model of gene action revealed a primarily additive genetic basis for floral characters associated with mating system variation. The exception was flower life span, which showed partial dominance of the outcrosser phenology. In contrast to similarly divergent species, the substantial differences in flower size between these A. uniflora populations appear to involve relatively few genes of large effect (minimum number of effective factors = 2.2 +/- 2.8 SE). In addition, correlations among traits in the F2 generation indicate that pleiotropy may be an important feature of the genetic architecture of floral evolution in A. uniflora. The evolution of selfing via major modifiers of floral morphology is consistent with other evidence for ecological selection for preemptive self-pollination in A. uniflora. Analyses of the genetic basis of autonomous selfing were complicated by hybrid breakdown in both F1 and F2 generations. Only F1 hybrids showed reductions in female fertility, but about 30% of F1 and F2 hybrids exhibited partial or complete male sterility. Male sterile flowers were characterized by short stamens, reduced petals, and a lack of protandry, as well as indehiscent anthers. This morphological breakdown mimics environmental disruptions of floral development and may result from novel genic interactions in hybrids.