昆虫孤雌生殖起源的遗传机制和进化意义

孤雌生殖在昆虫纲中有很高的发生比例,昆虫的孤雌生殖能力是造成农林灾害的一个重要而常被忽视的因素.孤雌生殖物种的存在和维持在进化生物学领域是个令人困惑的话题.在一些物种中孤雌生殖的细胞学机制得到了研究和解释,但是对于从两性生殖转换到孤雌生殖的遗传机制却知之甚少.了解孤雌生殖起源的遗传机制和进化意义对于预防农林虫害有着重要的意义.因此,该文就近年来对昆虫孤雌生殖起源的遗传和进化意义的研究做以综述,并就可能的利用前景进行简单讨论.     

孤雌生殖的爬行动物

无脊椎动物的不少门类中,如海绵动物、轮虫、线虫、环节动物和部分昆虫,具有孤雌生殖(Parthenogenesis)现象,这是大家熟知的。至于脊椎动物,已知无羊膜的鱼类和两栖动物中极少数物种有全雌居群,但其生殖方式则是雌核发生(Gynogenesis)。而在爬行动物这样的高等羊膜动物中确认存在着孤雌生殖的居群和物种,确是十分有趣的生物学问题。1958年苏联学者达莱夫斯基(I.S.Darevsky)首次报道了分布于亚美尼亚地区孤雌生殖的蜥蜴     

鱼类的天然雌核发育

1.孤雌生殖,雌核发育和杂合发育单性型种群在脊椎动物中是罕见的。尽管如此,到目前为止几乎已在各类脊椎动物中发现了单性型种群。该种群通常涉及3种繁殖方式,即孤雌生殖(parthenogenesis)、雌核发育(gynogenesis)和杂合发育(hybridogenesis)。     

钩盲蛇孤雌生殖的再证实

孤雌生殖是一种特殊的繁殖方式。行这种繁殖方式的动物,其卵细胞不需与精子发生受精作用即可开始卵裂及发育。换句话说,行孤雌生殖的动物,其雌性单方面就可繁衍后代。孤雌生殖的后代只承受了来自母体的染色体,它们也都是雌性,由之继续行孤雌生殖。当然,并不是说这些种类绝对没有雄性个体。其中某些种类在其分布的某些地区也曾发现少数雄性动物。孤雌生殖在较低等动物中是较普遍的一种繁殖方式,尤其在无脊椎动物中较为多见。在脊椎动物中,少数鱼类行孤雌生殖。爬行动物中,过去只知道一些蜥蜴行孤雌生殖。如壁虎科中的少数种类,其中包括产于我…     

脊椎动物基因组与基因复制研究进展

脊椎动物的出现是动物进化历史上一次质的飞跃.由于所有的脊椎动物在其胚胎发育中都呈现连续的解剖学特征,因此过去很多学者都根据现存脊椎动物的形态特征和在其发育过程中的解剖学特征假想原始脊椎动物,并推导其进化过程和起源.近年来的研究表明,通过对脊椎动物和与之亲缘关系接近的物种之间进行基因家族、染色体结构分析,可以对脊椎动物进化提供很多线索和证据.更多的研究表明,脊椎动物在进化过程中很可能发生过整体基因组的复制, 基因和/或基因组的复制可能是引起脊椎动物形体结构复杂性增加的根本原因.因此,基因和基因组的复制正在成为生物进化研究的热点问题.但这两种复制方式中哪一种是产生动物形体结构和功能复杂性增加最重要的原因尚有争论.     

鱼类的繁殖类型

鱼类与高等脊椎动物相比其繁殖方式具有多样性,不仅有卵生、胎生和卵胎生,还有孤雌生殖、雌核发育和杂合发育。     

哺乳动物卵的激活与孤雌发育

哺乳动物卵的激活与孤雌发育范必勤,邓满齐（江苏省农业科学院,南京,２１００１４）自然界孤雌生殖是一种繁殖方式,在无脊椎动物较为普遍,脊椎动物中除哺乳类外均有孤雌生殖现象［１］。孤雌生殖（Ｐａｒｔｈｅｎｏｓｅｎｅｓｉｓ）一词最早由Ｏｗｅｎ（１８４９）提出,系指没有雄性参与,由单个卵细胞产生个体的繁殖［２］。新近重新定义为无雄性配子的任何作用,由雌性配子产生的胚胎,不论其是否发育成个体,称为孤雌生殖。     

文昌鱼抗金葡菌感染差减文库的构建及免疫相关基因分析

适应性免疫的起源一直是免疫学研究的关键问题.文昌鱼被认为是最接近于脊椎动物的祖先 自从被发现以来一直是研究脊椎动物起源与进化机制的经典模式动物.为了在文昌鱼中寻找适应性免疫系统的分子证据,采用金黄色葡萄球菌感染文昌鱼以调查免疫的起源.应用抑制性差减杂交(SSH)技术,通过对差减文库克隆序列的测定,共获得588个表达序列标签(EST).对这些EST进行生物信息学分析和进一步功能分类,发现了一些免疫上调基因,如免疫调控基因、凋亡相关基因、细胞黏附相关基因、转录相关基因、信号传导相关基因等,以及一些非免疫相关基因;这些基因在文昌鱼中绝大多数为首次报道.金黄色葡萄球菌差减文库的成功构建,为调查文昌鱼抗细菌感染的分子事件提供了重要线索,对于这些新发现基因的进一步研究将有助于深入了解免疫系统起源与进化的机制.     

硬骨鱼中vasa基因的研究

vasa基因编码一个DEAD-box家族的ATP依赖性RNA解旋酶,最早在果蝇中发现.在绝大多数的脊椎动物和非脊椎动物中,vasa基因的杂交信号仅在生殖细胞系中特异性表达,因此它可以作为一种分子标签,广泛用于性腺发生、配子发生、原生殖细胞(primordial germ cells,PGCs)的起源、迁移、分化等方面的研究.综述了国内外有关硬骨鱼类vasa基因结构,表达与功能等方面的研究报道,并对其应用前景作了展望.     

单性脊椎动物的生殖方式

单性脊椎动物具有三种不同的生殖方式,即孤雌生殖、雌核生殖以及杂种生殖。本文比较了这三种生殖方式的异同,概述了各自配子发生的细胞学机制。     

DNA EVIDENCE FOR NONHYBRID ORIGINS OF PARTHENOGENESIS IN NATURAL POPULATIONS OF VERTEBRATES

Naturally occurring unisexual reproduction has been documented in less than 0.1% of all vertebrate species. Among vertebrates, true parthenogenesis is known only in squamate reptiles. In all vertebrate cases that have been carefully studied, the clonal or hemiclonal taxa have originated through hybridization between closely related sexual species. In contrast, parthenogenetic reproduction has arisen in invertebrates by a variety of mechanisms, including likely cases of “spontaneous” (nonhybrid) origin, a situation not currently documented in natural populations of vertebrates. Here, we present molecular data from the Neotropical night lizard genus Lepidophyma that provides evidence of independent nonhybrid origins for diploid unisexual populations of two species from Costa Rica and Panama. Our mitochondrial and nuclear phylogenies are congruent with respect to the unisexual taxa. Based on 14 microsatellite loci, heterozygosity (expected from a hybrid origin) is low in Lepidophyma reticulatum and completely absent in unisexual L. flavimaculatum. The unique value of this system will allow direct comparative studies between parthenogenetic and sexual lineages in vertebrates, with an enormous potential for this species to be a model system for understanding the mechanisms of nonhybrid parthenogenesis.     

Intra- and interspecific polymorphism of (AAT) n in microsatellite locus Du47D in parthenogenetic species of the genus Darevskia

The molecular structure of the allelic variants of (AAT) _n of the Du47D microsatellite locus was determined in parthenogenetic lizards Darevskia dahli, D. armeniaca, and D. rostombekovi. Comparative analysis of these alleles showed that they were characterized by perfect structure of microsatellite cluster, and were different in the number of (AAT) monomeric units, as well as in the combinations of species-specific substitutions and deletions in the microsatellite flanking regions. Molecular structure of microsatellite cluster, species-specific single nucleotide polymorphism (SNP), and different representation of alleles Du47 in the samples of parthenogenetic species examined point to the origin of the alleles from different bisexual species, which is consistent with the hybrid nature of unisexual species of the genus Darevskia. In addition, these data reflect different combination patterns of interspecific hybridization events with the participation of the same bisexual species upon the formation of hybrid genomes of parthenogenetic species. Possible application of the allelic variants of microsatellite loci of parthenogenetic lizards as the genetic markers for the analysis of the genomes of parthenogenetic species in the light of evolution, ecology, and parthenogenetic type of reproduction in vertebrates is discussed.     

ELECTROPHORETIC VARIATION IN THE PARTHENOGENETIC GRASSHOPPER WARRAMABA VIRGO AND ITS SEXUAL RELATIVES

Allozyme variation was examined within and between parthenogenetic clones of Warramabo virgo and the sexual ancestors, undescribed species P196 and P169. Both sexual species can be separated into northern and southern races using six loci, and the separate hybrid origin for the two major groups of parthenogenetic clones (the Standard Phylad and the Boulder-Zanthus Phylad) is substantiated by the racial variation in the sexual ancestors. Heterozygosity values in the parthenogenetic species are 6–9 times higher than those in the sexual species, and there is evidence for the accumulation of new variation subsequent to the hybrid origin of both phylads. The new variation is the result of either new mutations, recombination, or both. Three loci in the Standard Phylad clones reveal “orphan” alleles not found in the sexual ancestors; these alleles probably arose subsequent to hybridization but prior to the dispersal of the parthenogenetic clones. These data, in combination with those from other genetic studies, suggest that new variation may arise as a consequence of hybridization. Collectively, the allozyme, chromosome, molecular, and morphological data suggest that the Standard Phylad clones are of a more ancient but restricted origin, with clonal variation being the result of multiple hybridizations between individuals of P196 and P169.     

Genetic polymorphism and evolution in parthenogenetic animals. I. Polyploid curculionidae

The genetic variability at enzyme loci in different triploid and tetraploid parthenogenetic weevil populations has been elucidated by starch gel electrophoresis. The overall genotype of individual weevils belonging to different populations has been determined for over 25 loci. The results are compared with those obtained for diploid bisexual races of either the same or closely related species. The variation within a parthenogenetic population differs from that in diploid, sexually reproducing populations, i.e. the allele frequencies are not in a Hardy-Weinberg equilibrium. The results indicate that apomictic parthenogenetic populations can differentiate genetically. The genotypes within a population resemble each other more than genotypes belonging to different populations. It is evident that evolution still continues—even if slowed down—in parthenogenetic weevils. A comparison between the allele relationships in geographically isolated polyploid parthenogenetic populations and related diploid bisexual forms does not support the hypothetical hybrid origin of parthenogenesis and polyploidy in weevils. Parthenogenesis within a parthenogenetic weevil species is evidently monophyletic.     

Increased capacity for sustained locomotion at low temperature in parthenogenetic geckos of hybrid origin

The evolution of parthenogenesis is typically associated with hybridization and polyploidy. These correlates of parthenogenesis may have important physiological consequences that need be taken into account in understanding the relative merits of sexual and parthenogenetic reproduction. We compared the thermal sensitivity of aerobically sustained locomotion in hybrid/triploid parthenogenetic races of the gecko Heteronotia binoei and their diploid sexual progenitors. Endurance times at low temperature (10 degrees , 12.5 degrees , and 15 degrees C, 0.05 km h(-1)) were significantly greater in parthenogenetic females than in sexual females. Comparison of oxygen consumption rates during sustained locomotion at increasing speeds (0.05, 0.10, 0.15, 0.20, 0.25, and 0.30 km h(-1), 25 degrees C) indicated that parthenogenetic lizards have higher maximum oxygen consumption rates and maximum aerobic speeds than do female sexual geckos. In addition, parthenogenetic geckos showed greater levels of voluntary activity at 15 degrees C than did sexual geckos, although this pattern appears strongest in comparison to male sexual forms. Parthenogenetic lineages of Heteronotia thus have an advantage over sexual lineages in being capable of greater aerobic activity. This result is opposite of that found in prior studies of parthenogenetic teiid lizards (genus Cnemidophorus) and highlights the idiosyncratic nature of phenotypic evolution in parthenogens of hybrid origin.     

Functional rare males in diploid parthenogenetic Artemia

Functional males that are produced occasionally in some asexual taxa – called ‘rare males’ – raise considerable evolutionary interest, as they might be involved in the origin of new parthenogenetic lineages. Diploid parthenogenetic Artemia produce rare males, which may retain the ability to mate with females of related sexual lineages. Here, we (i) describe the frequency of male progeny in populations of diploid parthenogenetic Artemia, (ii) characterize rare males morphologically, (iii) assess their reproductive role, using cross‐mating experiments with sexual females of related species from Central Asia and characterize the F1 hybrid offspring viability and (iv) confirm genetically both the identity and functionality of rare males using DNA barcoding and microsatellite loci. Our result suggests that these males may have an evolutionary role through genetic exchange with related sexual species and that diploid parthenogenetic Artemia is a good model system to investigate the evolutionary transitions between sexual species and parthenogenetic strains.     

Multiple origins and rapid evolution of duplicated mitochondrial genes in parthenogenetic geckos (Heteronotia binoei; Squamata, Gekkonidae)

Accumulating evidence for alternative gene orders demonstrates that vertebrate mitochondrial genomes are more evolutionarily dynamic than previously thought. Several lineages of parthenogenetic lizards contain large, tandem duplications that include rRNA, tRNA, and protein-coding genes, as well as the control region. Such duplications are hypothesized as intermediate stages in gene rearrangement, but the early stages of their evolution have not been previously studied. To better understand the evolutionary dynamics of duplicated segments of mitochondrial DNA, we sequenced 10 mitochondrial genomes from recently formed ( approximately 300,000 years ago) hybrid parthenogenetic geckos of the Heteronotia binoei complex and 1 from a sexual form. These genomes included some with an arrangement typical of vertebrates and others with tandem duplications varying in size from 5.7 to 9.4 kb, each with different gene contents and duplication endpoints. These results, together with phylogenetic analyses, indicate independent and frequent origins of the duplications. Small, direct repeats at the duplication endpoints imply slipped-strand error as a mechanism generating the duplications as opposed to a false initiation/termination of DNA replication mechanism that has been invoked to explain duplications in other lizard mitochondrial systems. Despite their recent origin, there is evidence for nonfunctionalization of genes due primarily to deletions, and the observed pattern of gene disruption supports the duplication-deletion model for rearrangement of mtDNA gene order. Conversely, the accumulation of mutations between these recent duplicates provides no evidence for gene conversion, as has been reported in some other systems. These results demonstrate that, despite their long-term stasis in gene content and arrangement in some lineages, vertebrate mitochondrial genomes can be evolutionary dynamic even at short timescales.     

Phylogenetic evidence for hybrid origins of asexual lineages in an aphid species

Understanding the mode of origin of asexuality is central to ongoing debates concerning the evolution and maintenance of sexual reproduction in eukaryotes. This is because it has profound consequences for patterns of genetic diversity and ecological adaptability of asexual lineages, hence on the outcome of competition with sexual relatives both in short and longer terms. Among the possible routes to asexuality, hybridization is a very common mechanism in animals and plants. Aphids present frequent transitions from their ancestral reproductive mode (cyclical parthenogenesis) to permanent asexuality, but the mode of origin of asexual lineages is generally not known because it has never been thoroughly investigated with appropriate molecular tools. Rhopalosiphum padi is an aphid species with coexisting sexual (cyclically parthenogenetic) and asexual (obligately parthenogenetic) lineages that are genetically distinct. Previous studies have shown that asexual lineages of R. padi are heterozygous at most nuclear loci, suggesting either that they have undergone long-term asexuality (under which heterozygosity tends to increase) or that they have hybrid origins. To discriminate between these alternatives, we conducted an extensive molecular survey combining the sequence analysis of alleles of two nuclear DNA markers and mitochondrial DNA haplotypes in sexual and asexual lineages of R. padi. Both nuclear and cytoplasmic markers clearly showed that many asexual lineages have hybrid origins, the first such demonstration in aphids. Our results also indicated that asexuals result from multiple events of hybridization between R. padi and an unknown sibling species, and are of recent origin (contradicting previous estimates that asexual R. padi lineages were of moderate longevity). This study constitutes another example that putatively ancient asexual lineages are actually of much more recent origin than previously thought. It also presents a robust approach for testing whether hybrid origin of asexuality is also a common phenomenon in aphids.     

Developmental success, stability, and plasticity in closely related parthenogenetic and sexual lizards (Heteronotia, Gekkonidae)

The developmental trajectory of an organism is influenced by the interaction between its genes and the environment in which it develops. For example, the phenotypic traits of a hatchling reptile can be influenced by the organism's genotype, by incubation temperature, and by genetically coded norms of reaction for thermally labile traits. The evolution of parthenogenesis provides a unique opportunity to explore such effects: a hybrid origin of this trait in vertebrates modifies important aspects of the genotype (e.g., heterozygosity, polyploidy) and may thus impact not only on the phenotype generally, but also on the ways in which incubation temperature affects expression of the phenotype. The scarcity of vertebrate parthenogenesis has been attributed to developmental disruptions, but previous work has rarely considered reaction norms of embryogenesis in this respect. We used closely related sexual and asexual races of the Australian gecko Heteronotia binoei, which include those with multiple origins of parthenogenesis, to explore the ways in which reproductive modes (sexual, asexual), incubation temperatures (24, 27, and 30 degrees C), and the interaction between these factors affected hatchling phenotypes. The hatchling traits we considered included incubation period, incidence of deformities, hatchling survivorship, body size and shape, scalation (including fluctuating asymmetry), locomotor performance, and growth rate. Developmental success was slightly reduced (higher proportion of abnormal offspring) in parthenogenetic lineages although there was no major difference in hatching success. Incubation temperature affected a suite of traits including incubation period, tail length, body mass relative to egg mass, labial scale counts, running speed, growth rate, and hatchling survival. Our data also reveal an interaction between reproductive modes and thermal regimes, with the phenotypic traits of parthenogenetic lizards less sensitive to incubation temperature than was the case for their sexual relatives. Thus, the evolution of asexual reproduction in this species complex has modified both mean hatchling viability and the norms of reaction linking hatchling phenotypes to incubation temperature. Discussions on the reasons why parthenogenetic organisms are scarce in nature should take into account interactive effects such as these; future work could usefully try to tease apart the roles of parthenogenesis, its hybrid origin (and thus effects on ploidy and heterozygosity, etc.), and clonal selection in generating these divergent embryonic responses.     

Rapid Screening of DNA Diversity Using Dot-Blot Technology and Allele-Specific Oligonucleotides: Maternity of Hybrids and Unisexual Clones of Hybrid Origin (Lizards,Cnemidophorus)

Allele-specific oligonucleotide probes, together with dot-blot methods, can provide rapid and inexpensive screening of DNA types in large samples of organisms. Here we demonstrate their use in: (1) determining types of mitochondrial DNA in hundreds of lizards from a dynamic hybrid zone; (2) discovering intraspecific geographic variation in genes; and (3) determining and verifying the maternal ancestry of unisexual, parthenogenetic lizards in clones of hybrid origin. These methods are broadly applicable in research involving rapid screening of DNA types in large samples of specimens for any gene with sequence data from which to design specific probes.