被子植物花的起源：假说和证据

达尔文的“令人讨厌之谜”,即被子植物的起源和早期演化,一直是植物系统学研究领域的热点。被子植物区别于其它植物类群的一个显著特征就是花,因此,解决被子植物的起源之谜很大程度上取决于对被子植物花器官起源的研究,对被子植物花器官的详尽研究已经在形态、解剖、古植物、形态发生、分子等方面积累了大量的证据,植物学家基于这些证据为被子植物花器官的起源提出了各种各样的解释。综述了迄今为止被子植物花器官起源的主要学说流派,如：真花学说、假花学说、生殖叶学说、生殖茎节学说、生花植物学说、新假花学说、古草本学说和ANITA学说等。根据研究手段和获得证据的方式。作者将被子植物花器官起源研究划分为5个阶段,并简要阐述了各个阶段的代表学说和主要研究特点。     

花对称性的研究进展

花对称性(floral symmetry)是被子植物花部结构的典型特性之一,主要有辐射对称和两侧对称两种形式。被子植物初始起源的花为辐射对称,而两侧对称的花则是由辐射对称的花演变而来。两侧对称的花部结构是被子植物进化过程中的一个关键的革新,被认为是物种形成和分化的关键推动力之一。近年来有关花对称性的形成和进化机制的研究在植物学科的不同领域均取得了长足的进展。本文综述了花对称性在发育生物学、传粉生物学、生殖生态学及分子生物学等方面的研究进展。两侧对称形成于被子植物花器官发育的起始阶段,随后贯穿整个花器官发育过程或者出现在花器官发育后期的不同阶段。花器官发育过程中一种或多种类型器官的败育以及特异性花器官结构的形成是两侧对称形成的主要原因。研究表明,在传粉过程的不同阶段,花对称性均会受到传粉昆虫介导的选择作用。相比辐射对称的花,两侧对称的花提高了特异性传粉者的选择作用,增加了花粉落置的精确性,进而确保了其生殖成功。花对称性的分子机理已经在多种双子叶植物中进行了深入的研究。现有的证据表明,CYC同源基因在花对称性的分子调控方面起着非常重要的作用。花对称性在被子植物进化过程中是如何起源,与其他花部构成之间是否协同作用,一些不符合一般模式的科属其花对称性的形成机制等都是今后要进一步研究的命题。     

生花植物概念简介及其名称的商榷

在上个世纪最后的 2 0年里 ,系统学家应用形态性状对种子植物进行了大量的分支分析。其结果显示灭绝的五柱木属加上灭绝的本内苏铁目及尚存的买麻藤目是被子植物的姊妹群 ,形成一个强支 ,称之为生花植物支。生花植物假说对探讨被子植物起源有着重要影响 ,它激发人们讨论被子植物起源时间可能要提前到三叠纪甚至石炭纪 ,除了支持原有的真花学说外 ,还提出新恩格勒学说。但是 ,近年来对现存种子植物进行分子系统学研究的结果是 :(1)拒绝接受生花植物概念 ;(2 )买麻藤目并不是被子植物的姊妹群而是松柏目的姊妹群 ,甚至网结于松柏类而成为松科的姊妹群。这些结果并不使人惊讶 ,因为对探讨像包含许多灭绝类群的种子植物系统 ,决不可能是仅仅单独应用现代类群资料所能完成的。假如生花植物支是成立的 ,但其名称以AGPB支代替生花植物支可能较为合理。     

生花植物概念简介及其名称的商榷

在上个世纪最后的20年里,系统学家应用形态性状对种子植物进行了大量的分支分析。其结果显示灭绝的五柱木属加上灭绝的本内苏铁目及尚存的买麻藤目是被子植物的姊妹群,形成一个强支,称之为生花植物支。生花植物假说对探讨被子植物起源有着重要影响,它激发人们讨论被子植物起源时间可能要提前到三叠纪甚至石炭纪,除了支持原有的真花学说外,还提出新恩格勒学说。但是,近年来对现存种子植物进行分子系统学研究的结果是：(1)拒绝接受生花植物概念;(2)买麻藤目并不是被子植物的姊妹群而是松柏目的姊妹群,甚至网结于松柏类而成为松科的姊妹群。这些结果并不使人惊讶,因为对探讨像包含许多灭绝类群的种子植物系统,决不可能是仅仅单独应用现代类群资料所能完成的。假如生花植物支是成立的,但其名称以AGPB支代替生花植物支可能较为合理。     

被子植物系统学中花发育研究的进展及对今后研究的思考

从花发育研究的方法、花发育与被子植物花部结构的多样性、花发育与被子植物的系统发育以及 花发育的分子遗传学等四个方面对近年来被子植物系统学中花发育研究的主要进展作一综述,例举了 一些重要结果。同时,对该领域今后研究的方向和应注意的一些问题作了简要评论。作者认为植物的 形态结构可以看作是一个时空过程,在系统学研究中对花部性状的分析和认识应该树立动态的观点。 今后应该从动态的角度开展被子植物花的发生和发育以及性状在不同类群间的比较等方面的广泛研究,并加强对在被子植物花的起源和演化中起重要作用的花部同源异型现象的发育过程的观察。     

论三白草科的系统演化和地理分布

根据细胞学、孢粉学、花器官发生和发育、花部维管结构、胚胎学、比较形态及解剖学等方面的证据,对三白草科４属６种的系统演化关系进行了详细论证。并联系古地理、古气候和古植物的有关史料对三白草科分布区的起源作了据理推论。三白草科大约起源于早白垩纪古北大陆东南部。它的原始类型可能与现代三白草属的形态大体相似,为一种多年生具根状茎的草本,花小、无被、具苞片。白垩纪末期三白草的原始类群已完成它在古北大陆的迁移和     

被子植物花器官发育的模型演变和分子调控

基于模式植物拟南芥(Arabidopsis thaliana)和金鱼草(Antirrhinum majus)花器官突变体研究提出的四聚体模型揭示了花同源异型蛋白的相互作用方式;进一步提出的核小体拟态模型,解释了花同源蛋白四聚体调控目标靶基因的分子机理。被子植物花器官形态多样化与MADS-box基因的表达模式和功能分化密切相关。多年生被子植物花发育的高通量转录组分析表明,多种基因参与调控花器官发育过程。本文重点综述了被子植物花器官发育的模型演变、MADS-box基因结构和基因重复、miRNA调控以及相关转录组分析的最新研究成果,并对花器官发育的研究前景进行了展望。     

观察心皮叶性来源的好材料—日本晚樱

花是被子植物繁衍后代所特有的繁殖器官,由于花的各部分不易受外界环境的影响,其形态特征较稳定,变异较小,因此长期以来,人们都以花的形态结构作为被子植物分类鉴定和系统演化的主要依据。关于花的本质,目前普遍认为是枝性起源,即花像是一个变态的枝条,它的节间极度缩短,顶     

睡莲科系统位置评述

形态学研究显示睡莲科Nymphaeaceae具有许多原始性状。睡莲科又被称为“古草本”。最新的分子系统发育研究显示,睡莲科是现存被子植物系统树根部附近最早分异谱系的演化支之一,对研究被子植物(有花植物)的起源与早期进化具有重要价值,但有关睡莲科的范围和系统位置存在争议。被子植物的起源与辐射一直是植物学家关注的热点。本文对该科系统位置的研究历史与现状进行了评述。     

介绍学科动态,交流科研成果——全国植物形态解剖学学术讨论会记事

中国植物学会植物形态学专业委员会和浙江省植物学会联合主持召开的全国植物形态解剖学学术讨论会,于1991年11月2日至6日在杭州大学举行。参加会议的代表约120名。大会安排了十一个综述报告:(功植物解剖学的研究新进展,(2)中国植物形态解剖学的发展概况,(3)从顶枝学说、中柱学说、子叶节区学说看维管植物的形态演化和起源,(4)植物生态解剖学的研究进展,(,)被子植物种子形态解剖的研究进展,     

被子植物起源和早期演化研究的回顾与展望

近年来,被子植物起源和早期演化研究,由于手段和技术的更新,资料大量积累,取得了许多重要进展,成为植物学领域的一大热点。本文对过去近五十年的研究作了回顾,并从分子系统学、分支系统学、花原基发生的形态学、花发育的分子遗传学及白垩纪花和其它生殖结构化石研究等五个方面对该领域在最近十几年的研究进展进行综述,最后,对今后如何开展这方面的工作作了简要评论。     

Evolution of the angiosperms: calibrating the family tree.

Growing evidence of morphological diversity in angiosperm flowers, seeds and pollen from the mid Cretaceous and the presence of derived lineages from increasingly older geological deposits both imply that the timing of early angiosperm cladogenesis is older than fossil-based estimates have indicated. An alternative to fossils for calibrating the phylogeny comes from divergence in DNA sequence data. Here, angiosperm divergence times are estimated using non-parametric rate smoothing and a three-gene dataset covering ca. 75% of all angiosperm families recognized in recent classifications. The results provide an initial hypothesis of angiosperm diversification times. Using an internal calibration point, an independent evaluation of angiosperm and eudicot origins is performed. The origin of the crown group of extant angiosperms is indicated to be Early to Middle Jurassic (179-158 Myr), and the origin of eudicots is resolved as Late Jurassic to mid Cretaceous (147-131 Myr). Both estimates, despite a conservative calibration point, are older than current fossil-based estimates.     

Floral structure and evolution of primitive angiosperms: Recent advances

Concepts of primitive angiosperm flowers have changed in recent years due to new studies on relic archaic groups, new paleobotanical finds and the addition of molecular biological techniques to the study of angiosperm systematics and evolution.Magnoliidae are still the hot group, but emphasis is now on small primitive flowers with few organs and also on the great lability of organ number. Of the extant groups, a potential basal position of the paleoherbs has been discussed by some authors. Although some paleoherbs have a simple gynoecium with a single orthotropous ovule, anatropous ovules may still be seen as plesiomorphic in angiosperms. Anatropy is not necessarily a consequence of the advent of closed carpels. It may also exhibit biological advantages under other circumstances as is the case in podocarps among gymnosperms. Valvate anthers have now been found in most larger subgroups of theMagnoliidae (recently also in paleoherbs) and in some Cretaceous fossils. Nevertheless, as seen from its systematic distribution, valvate dehiscence is not necessarily plesiomorphic for the angiosperms, but may be a facultative by-product of the thick connectives and comparatively undifferentiated anther shape inMagnoliidae and lowerHamamelididae. A perianth is relatively simple in extantMagnoliidae or even wanting in some families. In groups with naked flowers the perianth may have been easily lost because integration in the floral architecture was less pronounced than in more advanced angiosperm groups. Problems with the comparison of paleoherb flowers with those ofGnetales are discussed. The rapid growth of information from paleobotany and molecular systematics requires an especially open attitude towards the evaluation of various hypotheses on early flower evolution in the coming years.     

Evolution of sex: role of deleterious mutation and mobile elements

Prevalence of sexual reproduction is still enigma. The main character of sex is alleles mixing that could be advantageous either in unstable environment (in this case sex provides high temp of evolution) or in unstable genotype (in this case sex provides purge of genome from deleterious mutations). As long as not all species inhabit highly changeable environments, variation of genotypes is more important factor. As the majority of new mutations is deleterious, effective mechanism of genome purging is needed. Maintenance of "purging mechanism" may be a single role of sex. Two promising mutational hypotheses--clade selection (Muller's ratchet and Nunney's hypothesis) and mutational deterministic hypothesis of Kondrashov claim that more effective elimination of slightly-deleterious mutations provides main advantage to sexual population in comparison with asexual. Despite prima facie similarity, these hypotheses differ in mechanisms, work at different temporal scales and have different consequences. Kondrashov's hypothesis reveals short-term advantage of sexual reproduction, and thus, based on the individual selection. Clade selection displays long-term advantage of sexual reproduction that could be realized only by group selection. The role of mobile elements in evolution of sexual reproduction is also discussed. Firstly, mobile elements ("sexual molecular parasites") can complicate the problem: having been domesticated in asexual genomes and remaining active in sexual genomes they lead to higher mutational rate in sexual organisms and so violate assumption critical for both mutational hypotheses of "other things being equal". Secondly, mobile elements could be leader factor of origin of sex (hypothesis proposed by Hickey). Because theory of group selection could explain maintenance of sex, but not its origin, mobile elements could induce the origin of sex but were not able to maintain it, so the next scenario of evolution of sex is proposed: mobile elements induced origin of sex, which was established later by group selection because provided long term benefit (Muller's ratchet and Nunney's hypothesis). So, on all stages of evolution, sex was not advantageous for the organism per se.     

Life history, floral development, and mating system in Clarkia xantiana (Onagraceae): do floral and whole-plant rates of development evolve independently?

Autogamously self-fertilizing taxa have evolved from outcrossing progenitors at least 12 times in the annual wildflower genus, Clarkia (Onagraceae). In C. xantiana, individuals of the selfing subspecies (ssp. parviflora) flower at an earlier age, produce successive flowers more rapidly, and produce flowers that complete their development more rapidly than their outcrossing counterparts (ssp. xantiana). Two hypotheses have been proposed to explain the joint evolution of these whole-plant and individual floral traits. The accelerated life cycle hypothesis proposes that selection favoring a short life cycle in environments with short growing seasons (such as those typically occupied by parviflora) has independently favored genotypes with early reproduction, synchronous flower production, and rapidly developing, self-fertilizing flowers. The correlated response to selection hypothesis similarly proposes that selection in environments with short growing seasons favors early reproduction, but that rapid floral development and increased selfing evolve as correlated responses to selection due to genetic linkage (or pleiotropy) affecting both whole-plant and floral development. We conducted a greenhouse experiment using maternal families from two field populations of each subspecies to examine covariation between floral and whole-plant traits within and among populations to seek support for either of these hypotheses. Our results are consistent with the accelerated life cycle hypothesis but not with the correlated response to selection hypothesis.     

Intralocus sexual conflict can drive the evolution of genomic imprinting

Genomic imprinting is a phenomenon whereby the expression of an allele differs depending upon its parent of origin. There is an increasing number of examples of this form of epigenetic inheritance across a wide range of taxa, and imprinting errors have also been implicated in several human diseases. Various hypotheses have been put forward to explain the evolution of genomic imprinting, but there is not yet a widely accepted general hypothesis for the variety of imprinting patterns observed. Here a new evolutionary hypothesis, based on intralocus sexual conflict, is proposed. This hypothesis provides a potential explanation for much of the currently available empirical data, and it also makes new predictions about patterns of genomic imprinting that are expected to evolve but that have not, as of yet, been looked for in nature. This theory also provides a potential mechanism for the resolution of intralocus sexual conflict in sexually selected traits and a novel pathway for the evolution of sexual dimorphism.     

The evolutionary origin of the durophagous pelagic stingray ecomorph

Studies of the origin of evolutionary novelties (novel traits, feeding modes, behaviours, ecological niches, etc.) have considered a number of taxa experimenting with new body plans, allowing them to occupy new habitats and exploit new trophic resources. In the marine realm, colonization of pelagic environments by marine fishes occurred recurrently through time. Stingrays (Myliobatiformes) are a diverse clade of batoid fishes commonly known to possess venomous tail stings. Current hypotheses suggest that stingrays experimented with a transition from a benthic to a pelagic/benthopelagic habitat coupled with a transition from a non-durophagous diet to extreme durophagy. However, there is no study detailing macroevolutionary patterns to understand how and when habitat shift and feeding specialization arose along their evolutionary history. A new exquisitely preserved fossil stingray from the Eocene Konservat-Lagerstätte of Bolca (Italy) exhibits a unique mosaic of plesiomorphic features of the rajobenthic ecomorph, and derived traits of aquilopelagic taxa, that helps to clarify the evolutionary origin of durophagy and pelagic lifestyle in stingrays. A scenario of early evolution of the aquilopelagic ecomorph is proposed based on new data, and the possible adaptive meaning of the observed evolutionary changes is discussed. The body plan of †Dasyomyliobatis thomyorkei gen. et sp. nov. is intermediate between the rajobenthic and more derived aquilopelagic stingrays, supporting its stem phylogenetic position and the hypothesis that the aquilopelagic body plan arose in association with the evolution of durophagy and pelagic lifestyle from a benthic, soft-prey feeder ancestor.     

Ancestral xerophobia: a hypothesis on the whole plant ecophysiology of early angiosperms

Today, angiosperms are fundamental players in the diversity and biogeochemical functioning of the planet. Yet despite the omnipresence of angiosperms in today's ecosystems, the basic evolutionary understanding of how the earliest angiosperms functioned remains unknown. Here we synthesize ecophysiological, paleobotanical, paleoecological, and phylogenetic lines of evidence about early angiosperms and their environments. In doing so, we arrive at a hypothesis that early angiosperms evolved in evermoist tropical terrestrial habitats, where three of their emblematic innovations – including net‐veined leaves, xylem vessels, and flowers – found ecophysiological advantages. However, the adaptation of early angiosperm ecophysiology to wet habitats did not initially promote massive diversification and ecological dominance. Instead, wet habitats were permissive for the ecological roothold of the clade, a critical phase of early diversification that entailed experimentation with a range of functional innovations in the leaves, wood, and flowers. Later, our results suggest that some of these innovations were co‐opted gradually for new roles in the evolution of greater productivity and drought tolerance, which are characteristics seen across the vast majority of derived and ecologically dominant angiosperms today.     

Mitochondrial genes from 18 angiosperms fill sampling gaps for phylogenomic inferences of the early diversification of flowering plants

The early diversification of angiosperms is thought to have been a rapid process, which may complicate phylogenetic analyses of early angiosperm relationships. Plastid and nuclear phylogenomic studies have raised several conflicting hypotheses regarding overall angiosperm phylogeny, but mitochondrial genomes have been largely ignored as a relevant source of information. Here we sequenced mitochondrial genomes from 18 angiosperms to fill taxon-sampling gaps in Austrobaileyales, magnoliids, Chloranthales, Ceratophyllales, and major lineages of eudicots and monocots. We assembled a data matrix of 38 mitochondrial genes from 107 taxa to assess how well mitochondrial genomic data address current uncertainties in angiosperm relationships. Although we recovered conflicting phylogenies based on different data sets and analytical methods, we also observed congruence regarding deep relationships of several major angiosperm lineages: Chloranthales were always inferred to be the sister group of Ceratophyllales, Austrobaileyales to mesangiosperms, and the unplaced Dilleniales was consistently resolved as the sister to superasterids. Substitutional saturation, GC compositional heterogeneity, and codon-usage bias are possible reasons for the noise/conflict that may impact phylogenetic reconstruction; and angiosperm mitochondrial genes may not be substantially affected by these factors. The third codon positions of the mitochondrial genes appear to contain more parsimony-informative sites than the first and second codon positions, and therefore produced better resolved phylogenetic relationships with generally strong support. The relationships among these major lineages remain incompletely resolved, perhaps as a result of the rapidity of early radiations. Nevertheless, data from mitochondrial genomes provide additional evidence and alternative hypotheses for exploring the early evolution and diversification of the angiosperms.     

Things that go bump in the night: evolutionary interactions between cephalopods and cetaceans in the tertiary

Echolocation has evolved independently in several vertebrate groups, and hypotheses about the origin of echolocation in these groups often invoke abiotic mechanisms driving morphological evolution. In bats, for example, the ecological setting associated with the origin of echolocation has been linked to global warming during the Palaeocene–Eocene; similarly, the origin of toothed whales (odontocetes) has been broadly correlated with the establishment of the circum-Antarctic current. These scenarios, and the adaptational hypotheses for the evolution of echolocation with which they are associated, neglect a consideration of possible biotic mechanisms. Here we propose that the origin of echolocation in odontocetes was initially an adaptation for nocturnal epipelagic feeding – primarily on diel migrating cephalopods. We test this hypothesis using data on the temporal, geographical, and water column distributions of odontocetes and cephalopods, and other global events from their respective tertiary histories. From this analysis, we suggest that echolocation in early odontocetes aided nocturnal feeding on cephalopods and other prey items, and that this early system was exapted for deep diving and hunting at depths below the photic zone where abundant cephalopod resources were available 24 h a day. This scenario extends to the evolution of other cephalopod feeding (teuthophagous) marine vertebrates such as pinnipeds and Mesozoic marine reptiles.