被子植物异型花柱及其进化意义

异型花柱(heterostyly)是被子植物中一种特殊的花多态现象和雌雄异位形式,包括二型花柱(distyly)和三型花柱(tristyly)2种类型.据报道,在被子植物的约31个科中有异型花柱植物.该类型植物的花部特征在避免自交、促进准确的异交传粉以及通过降低雌雄功能干扰以提高亲本适合度等方面具有重要的进化意义.该文从以下3个方面总结和分析了异型花柱及其进化意义:(1)异型花柱植物的类型和花部特征、附属多态性和种群结构:(2)异型花柱植物在被子植物中的分布、起源和演化:(3)异型花柱植物的进化适应意义.结合目前作者开展的有关工作,对异型花柱植物研究中存在的一些问题进行讨论和展望,希望能为国内工作者开展该领域的研究提供一些参考.     

被子植物镜像花柱及其进化意义

镜像花柱是指花柱在花水平面上向左(左花柱型)或向右(右花柱型)偏离花中轴线,是一种花柱多态现象,可根据左、右花柱花在植株上的排列式样划分为单型镜像花柱和二型镜像花柱两类,或根据镜像花柱和雄蕊的排列方式划分为雌雄互补镜像花柱和非雌雄互补镜像花柱两类。镜像花柱现象已在被子植物11个科的部分种类中进行了报道,它在保护功能型雄蕊和雌蕊、通过自交产生繁殖保障效应以及通过减小雌雄功能间干扰、提高异交率和雄性适合度等方面具有重要的进化意义,目前已成为植物繁殖生物学领域的研究热点。本文总结分析了国际上有关镜像花柱的研究工作,重点介绍:(1)镜像花柱的类型、镜像花的形态分化及花部特征,(2)镜像花柱植物在被子植物中的分布及其遗传演化,(3)镜像花的交配式样和交配频率、传粉特点及其进化生物学意义,并对今后的研究方向进行了展望,以期为进一步研究镜像花柱植物的进化生物学特性,推动我国在该领域的发展提供科学依据。     

水生被子植物的繁育系统与进化

对植物繁育系统的多样化及其作用模式和机制的研究,是理解植物各类群进化的一个重要基础。这一观点,已被从事植物进化生物学研究的国内外学者普遍接受。本世纪三十年代基因流概念的引人,促使人们在进化研究中,对植物繁育系统的重要性进行了重新评价。在过去的半个多世纪,植物进化生物学也因之而面貌一新,并得到了飞速的发展。时至今日,对植物进化过程中与繁育系统密切相关的一些重要机制,已有了较为深刻的认识（如有性过程中的遗传重组与植物的变异、进而与居群进化之间的关系等）。遗憾的是,这些认识几乎都是建立在对陆生植物研究…     

花柱卷曲性异交机制及其进化生态学意义

 有花植物具有纷繁复杂的繁育系统,以避免或促进自交或异交。花柱卷曲性异交机制(Flexistyly)是最近在热带山姜属(Alpinia)植物中发现的一种促进异交的行为机制。具有这一机制的种类其自然种群中的个体根据开花行为的不同分为两种表型：一种上午散发花粉而其柱头向上反卷,远离昆虫拜访的通道;另一种其柱头上午垂向唇瓣,能够接受拜访昆虫的传粉,但自身的花药却不打开。到中午时分,两种表型花通过互为相反的柱头卷曲运动转换性别——前者柱头向下卷曲,后者柱头向上卷曲且花药打开。每朵花的花期为12 h,两种表型在自然种     

艳丽耳草的二型花柱及异型自交不亲和系统

异型花柱是一种受遗传因素控制的花型多态性现象,包括二型花柱和三型花柱两种类型.本文以茜草科艳丽耳草(Hedyotis pulcherrima)为实验材料,通过对其野外居群的花型、花部形态及花粉特征等观察,发现艳丽耳草野外居群同时存在长花柱型花和短花柱型花,长/短花柱型花的数量比例为1∶1.两型花具有精确的交互式雌雄异位特征,并且该特征与花冠长度相关性显著.长/短花柱型花的柱头裂片长度、花粉大小及淀粉含量等具有二型性.花粉体外培养时花粉萌发率及花粉管生长速率无显著性差异.人工授粉后,艳丽耳草长/短花柱型花型间异交花粉管生长形态正常,授粉24 h后花粉管均已进入子房.而长/短花柱型花在自交及型内异交下均表现为不亲和,花粉管生长停止于柱头,花粉管顶端累积胼胝质并膨大.艳丽耳草没有无融合生殖现象,型间人工辅助异交授粉结实率为100％,显著高于自然结实率.本研究结果表明,艳丽耳草是典型的二型花柱植物,并具有异型自交不亲和系统.     

被子植物雌全同株性系统:系统演化、性表达与进化意义

雌全同株是指雌花和两性花共同发生在同一植株上的性表达形式。作为被子植物从雌雄同花(两性花)向雌雄同株异花进化的一个重要阶段,雌全同株性系统在减少昆虫对雌性的取食和伤害、提高异交率以减少近交衰退、减少雌/雄功能干扰、提高雌/雄性功能间资源分配的灵活性,以及吸引传粉者等方面具有重要的进化适应意义。根据APG III分类系统,雌全同株性系统在被子植物木兰分支(magnoliids)的短蕊花科、单子叶植物分支(monocots)的天南星科和禾本科,以及核心真双子叶植物分支(core eudicots)中的菊科、苋科、唇形科和石竹科等23科中均有报道,且以菊科植物中最多。雌全同株植物不同类群的雌花和两性花在位置、形态、大小及开花时间等性表达特征上表现出多样化,且这些特征不仅受遗传因子的调控,还受可获得资源(如营养、光照、温度和水分等条件)的制约。该文针对我国对雌全同株性系统的研究还相对较少的现状,重点对具雌全同株性系统的类群在被子植物中的分布与系统演化、性表达与环境的关系等方面进行了分析与总结,并对有关其进化适应意义的5个假说进行了介绍和评价,对今后的研究方向进行了展望,以期为推动我国对被子植物雌全同株性系统的进化式样与机制研究提供理论资料。     

被子植物在进化中与环境相适应的传粉机制

被子植物主要依靠环境中的各种媒介传粉才能受精,产生种子。如果没有传粉机制,被子植物将不能繁殖而最终灭亡。概述了被子植物在进化过程中出现的多种多样传粉机制,对繁殖与进化生物学的研究有重要意义。     

异型花柱植物喀什补血草的传粉者功能群与花粉转移效率

异型花柱是受遗传控制的花柱多态现象, 被达尔文认为是植物通过在传粉者体表不同部位滞落花粉以促进型间花粉准确传递的一种适应。该现象虽已受到广泛关注, 但在一些花型变异较大且不稳定的传粉系统中, 不同传粉者对各花型繁殖所产生的影响仍知之甚少。该研究以分布于新疆天山南坡的一个有同长花柱共存的异型花柱植物喀什补血草(Limonium kaschgaricum)种群为研究对象, 对其花型构成及频率、传粉者及花粉转移效率等进行了调查分析。结果表明: 1)种群中除了存在雌/雄蕊长度交互对应的长(L)/短(S)花柱型花外, 还有雌/雄蕊同长的花(H型), 且各花型花的花冠口直径、花冠筒长及花粉量等参数间无差异, 但花粉纹饰和柱头乳突细胞形态具二型性。其中, H型花的花粉和柱头形态与L型花(或S型花)的一致。2)花型内和自花授粉均不亲和; 型间授粉时, 花粉和柱头形态不同的花型间亲和, 反之不亲和。3)种群内存在长/短吻两类传粉昆虫。在以短吻传粉者为主的盛花初、中期, L和H型花柱头上的异型花粉数均显著高于S型花的, 且L和S型花高位性器官间的异型花粉传递效率高于低位性器官间的; 而在以长吻传粉者为主的盛花后期, L和S型花的柱头间异型花粉数无显著差异, 且高/低位性器官间具有相同的异型花粉转移效率; 与传粉者出现时期相对应的、在花期不同阶段开放花的结实率也明显不同。4)长/短吻昆虫具明显不同的传粉功能, 短吻昆虫只能对L和H型花进行有效传粉, 且访花频率和型间花粉转移效率较低, 为低效传粉者; 而长吻昆虫对各花型均能有效传粉, 具高的访花频率和型间花粉转移效率, 为高效传粉者。因为长吻昆虫的阶段性出现所形成的不稳定传粉系统, 使低效的短吻昆虫可能会成为种群中花型变异的驱动力, 并使S型花受到更大的选择压力。H型花克服了柱头缩入的弊端, 可能会成为不稳定传粉系统下的一个替代花型而持续存在。     

核rDNA的ITS序列在被子植物系统与进化研究中的应用

被子植物与其它高等真核生物相似,核rDNA是高度重复的串联序列。由于同步进化的力量,绝 大多数物种中这些重复单位间已发生纯合或接近纯合。核rDNA的内转录间隔区(ITS)包含被5.8S rDNA所分隔的ITS1和ITS2两个片段,ITSl的长度为187～298bp,ITS2为187～252bp,经PCR扩 增后可以方便地对这两个片段进行直接测序或克隆测序。ITS序列变异较快,可以提供较丰富的变异 位点和信息位点,已证实它是研究许多被子植物类群系统与进化的重要分子标记,不仅可用于解决科、 亚科、族、属、组内的系统发育和分类问题,而且可用于重建多倍体复合体的网状进化关系,探讨异源多倍体的起源过程,然而,正是由于ITS序列变异较快,它一般不适于科以上水平的系统发育研究。     

连翘的二型花柱

报道了连翘中的二型花柱(长花柱和短花柱)。长花柱的柱头和雄蕊高度分别为6.12±0.05 mm和2.35±0.04mm,短花柱则为2.23±0.04mm和6.02±0.06mm。短花柱花的花冠大小明显超过长花柱。开放授粉条件下,长短花柱花的座果率分别为9.11 %±0.04%和8.93 %±0.07%。人工异交的座果率在长-短(36.8%±0.04%)与短-长(36.2%±0.07%)组合间无明显差异(F(1,39)=1 .38,P=0.14)。人工异交实验表明,传粉者限制可能发生在生长于中国东北部的人工连翘种群中,这可能是因为该地区早春的低温和多风气候条件影响传粉者的种类和活动。     

The Rhinal Bone and Its Evolutionary Significance

Bjerring, H. C. (Section of Palaeozoology, Swedish Museum of Natural History, Stockholm, Sweden.) The rhinal bone and its evolutionary significance. Zool. Scripta 1 (5): 193–201, 1972.– On the basis of serially sectioned embryos of Amia calva, the ethmoidal region of the endocranium and certain adjacent exo-skeletal elements are analysed. The results include evidence of the existence of branchial moieties pertaining to the first or premandibular metamere. Each of these moieties comprises infrapharyngobranchial (the ethmobasal), suprapharyngobranchial (the orbitonasal lamina), and epibranchial (the palatoquadrate pterygoid process) endoskeletal components as well as horizontal infrapharyngeal (the vomer), ascending infrapharyngeal (the rhinal), and epal (the dermopalatines and ectopterygoid) dental plates. The anterior cerebral arteries may represent the efferent blood vessels of the first-metamere branchial moieties. A hypothesis on the origin of the nasal sacs is offered.      

The Genomic Selfing Syndrome Accompanies the Evolutionary Breakdown of Heterostyly

The evolutionary transition from outcrossing to selfing can have important genomic consequences. Decreased effective population size and the reduced efficacy of selection are predicted to play an important role in the molecular evolution of the genomes of selfing species. We investigated evidence for molecular signatures of the genomic selfing syndrome using 66 species of Primula including distylous (outcrossing) and derived homostylous (selfing) taxa. We complemented our comparative analysis with a microevolutionary study of P. chungensis, which is polymorphic for mating system and consists of both distylous and homostylous populations. We generated chloroplast and nuclear genomic data sets for distylous, homostylous, and distylous–homostylous species and identified patterns of nonsynonymous to synonymous divergence (d_N/d_S) and polymorphism (π_N/π_S) in species or lineages with contrasting mating systems. Our analysis of coding sequence divergence and polymorphism detected strongly reduced genetic diversity and heterozygosity, decreased efficacy of purifying selection, purging of large-effect deleterious mutations, and lower rates of adaptive evolution in samples from homostylous compared with distylous populations, consistent with theoretical expectations of the genomic selfing syndrome. Our results demonstrate that self-fertilization is a major driver of molecular evolutionary processes with genomic signatures of selfing evident in both old and relatively young homostylous populations.     

The Classification and Functional Significance of Staminodes in Angiosperms

All staminodes in an androecium fail to produce viable pollen grains and cannot contribute directly to male fitness. Staminodes are identified in the flowers of approximately 54% of known genera representing >32% of all angiosperm families. The functional morphology and biochemistry of staminodes differs significantly from stamens with fertile anthers. In the absence of sperm production, some staminodes evolved novel adaptations contributing to the reproductive success of their flowers. We subdivided these staminodes into eight functional types: 1) Staminodes offering visual/olfactory cues; 2) Staminodes offering edible/inedible rewards; 3) Staminodes that deceive pollinators with false rewards; 4) Staminodes facilitating or directing the movements of pollinators in flowers;5) Stami nodes that facilitate stigma movement; 6) Staminodes functioning as secondary pollen presenters; 7) Staminodes that protect other floral organs; 8) Staminodes that prevent mechanical self pollination (autogamy). As a component within a flower, the majority of staminodes function as promoters of reproductive success by interacting directly with the pollinator to increase pollination efficiency (both pollen dispersal and deposition). Therefore, it is not surprising that one staminode may have more than one function over the flower′s lifespan and is closely associated with the size, abundance, behavior and taxonomic diversity of pollinators, floral predators and thieves. To correctly evaluate the function of staminodes, multi disciplinary approach using a range of protocols, equipment and materials is suggested. This approach allows us to compare the roles different staminodes play in the reproductive success of both closely and distantly related angiosperms, then uncover their evolutionary significance in angiosperm diversification.     

Elasmobranch Central Nervous System Organization and Its Possible Evolutionary Significance

Examination of shark brain:body ratios reveals that these taxapossess relative brain volumes in a range overlapping thoseof bony fish as well as birds and mammals. Much of the variationis due to relative development of the telencephalon and cerebellum.Telencephalic weights vary from 24% in Squalus to 52% in Sphyrna.Analysis of the cytoarchitectonics of the shark brains revealsat least two patterns of development. Squalomorph sharks possesslow brain:body ratios, and the telencephalon of these taxa possesswell developed lateral ventricles and poorly developed pallialareas. The diencephalon is characterized by prominent periventricularlaminae, and the cerebellum lacks foliation. The lamniform andcarcharhiniform sharks are characterized by high brain: bodyratios, and there is marked hypertrophy of the telencephalon.The roof (pallial) regions, as well as the diencephalon, arecharacterized by extensive cellular migrations. The cerebellaof these forms possess extensive complex foliation. These brain patterns are compared with the brain organizationof Holocephali, and I conclude that the holocephalans are asister radiation of the elasmobranchs. Comparisons with bonyfish and land vertebrates suggest that elasmobranchs have independentlydeveloped complex pallial fields and cerebellar foliation asa result of parallel evolutionary trends.     

被子植物花粉形成过程中细胞骨架的动态及生物学意义

本文对被子植物花粉形成过程中细胞骨架(微管和微丝)的分布格局、动态变化及可能的功能进行了概括,并对相关的几个有争议的问题进行了讨论。     

被子植物花粉形成过程中细胞骨架的动态及生物学意义

本文对被子植物花粉形成过程中细胞骨架（微管和微丝）的分布格局、动态变化及可能的功能进行了概括,并对相关的几个有争议的问题进行了讨论     

Pollen Morphology of the Genus Malus and Its Taxonomic and Evolutionary Significance

The pollen morphology of 26 species and 5 hybrids of the genus Malus was investigated with aid of SEM. It is found that the pollen morphology of Malus is rather similar in shape, size, position and number of aperture and exine sculpture. The characters of polle morphology of sections and series are as follows: Sect. Malus Ser. Baccatae: Striae regular, parallel to colpi, and conjunct at pole; Ser. Pumilae: the same as in Ser. Baccatae, but more or less curved near pole. Sect. Docyniopes: Striae regular and parallel to colpi, but bent near pole. Sect. Chloromeles: Striae irregular, dense and interlock. Sect. Sorbormalus: Striae irregular, sparse and not interlock. Ser. Sieboldiance: pollen grains prolate, with perforation among striae, colpi narrow; Ser. Kansuenses: Pollen grains spheroidal, colpi wide in the middle but narrow at both ends, striae relatively dense and regular, mostly dichotomous, perforation present; Ser. Yunnanenses; Pollen grain spheroidae, colpi wide in the middle but narrow at both ends, striae obviously irregular, less dichotomous, perforation absent. The major evolutionary trend of exine sculpture of pollen may be from densely thin-striate to sparsely striate with perforation. Characters of the exine sculpture of hybrids can be used to recognize the rela-tionship between the parental species.     

Evolutionary Dynamics of the Leucine-Rich Repeat Receptor-Like Kinase (LRR-RLK) Subfamily in Angiosperms

Gene duplications are an important factor in plant evolution, and lineage-specific expanded (LSE) genes are of particular interest. Receptor-like kinases expanded massively in land plants, and leucine-rich repeat receptor-like kinases (LRR-RLK) constitute the largest receptor-like kinases family. Based on the phylogeny of 7,554 LRR-RLK genes from 31 fully sequenced flowering plant genomes, the complex evolutionary dynamics of this family was characterized in depth. We studied the involvement of selection during the expansion of this family among angiosperms. LRR-RLK subgroups harbor extremely contrasting rates of duplication, retention, or loss, and LSE copies are predominantly found in subgroups involved in environmental interactions. Expansion rates also differ significantly depending on the time when rounds of expansion or loss occurred on the angiosperm phylogenetic tree. Finally, using a d_N/d_S-based test in a phylogenetic framework, we searched for selection footprints on LSE and single-copy LRR-RLK genes. Selective constraint appeared to be globally relaxed at LSE genes, and codons under positive selection were detected in 50% of them. Moreover, the leucine-rich repeat domains, and specifically four amino acids in them, were found to be the main targets of positive selection. Here, we provide an extensive overview of the expansion and evolution of this very large gene family.Receptor-like kinases (RLKs) constitute one of the largest gene families in plants and expanded massively in land plants (Embryophyta; Lehti-Shiu et al., 2009, 2012). For plant RLK gene families, the functions of most members are often not known (especially in recently expanded families), but some described functions include innate immunity (Albert et al., 2010), pathogen response (Dodds and Rathjen, 2010), abiotic stress (Yang et al., 2010), development (De Smet et al., 2009), and sometimes multiple functions (Lehti-Shiu et al., 2012). The RLKs usually consist of three domains: an N-terminal extracellular domain, a transmembrane domain, and a C-terminal kinase domain (KD). In plants, the KD usually has a Ser/Thr specificity (Shiu and Bleecker, 2001), but Tyr-specific RLKs were also described (e.g. BRASSINOSTEROID INSENSITIVE1; Oh et al., 2009). Interestingly, it was estimated that approximately 20% of RLKs contain a catalytically inactive KD (e.g. STRUBBELIG and CORYNE; Chevalier et al., 2005; Castells and Casacuberta, 2007; Gish and Clark, 2011). In Arabidopsis (Arabidopsis thaliana), 44 RLK subgroups (SGs) were defined by inferring the phylogenetic relationships between the KDs (Shiu and Bleecker, 2001). Interestingly, different SGs show different duplication/retention rates (Lehti-Shiu et al., 2009). Specifically, RLKs involved in stress responses show a high number of tandemly duplicated genes whereas those involved in development do not (Shiu et al., 2004), which suggests that some RLK genes are important for the responses of land plants to a changing environment (Lehti-Shiu et al., 2012). There seem to be relatively few RLK pseudogenes compared with other large gene families, and copy retention was argued to be driven by both drift and selection (Zou et al., 2009; Lehti-Shiu et al., 2012). As most SGs are relatively old and RLK subfamilies expanded independently in several plant lineages, duplicate retention cannot be explained by drift alone, and natural selection is expected to be an important driving factor in RLK gene family retention (Lehti-Shiu et al., 2009).Leucine-rich repeat-receptor-like kinases (LRR-RLKs), which contain up to 30 leucine-rich repeat (LRRs) in their extracellular domain, constitute the largest RLK family (Shiu and Bleecker, 2001). Based on the KD, 15 LRR-RLK SGs have been established in Arabidopsis (Shiu et al., 2004; Lehti-Shiu et al., 2009). So far, two major functions have been attributed to them: defense against pathogens and development (Tang et al., 2010b). LRR-RLKs involved in defense are predominantly found in lineage-specific expanded (LSE) gene clusters, whereas LRR-RLKs involved in development are mostly found in nonexpanded groups (Tang et al., 2010b). It was also discovered that the LRR domains are significantly less conserved than the remaining domains of the LRR-RLK genes (Tang et al., 2010b). In addition, a study of four plant genomes (Arabidopsis, grape [Vitis vinifera], poplar [Populus trichocarpa], and rice [Oryza sativa]) showed that LRR-RLK genes from LSE gene clusters show significantly more indications of positive selection or relaxed constraint than LRR-RLKs from nonexpanded groups (Tang et al., 2010b).The genomes of flowering plants (angiosperms) have been shown to be highly dynamic compared with most other groups of land plants (Leitch and Leitch, 2012). This dynamic is mostly caused by the frequent multiplication of genetic material, followed by a complex pattern of differential losses (i.e. the fragmentation process) and chromosomal rearrangements (Langham et al., 2004; Leitch and Leitch, 2012). Most angiosperm genomes sequenced so far show evidence for at least one whole-genome multiplication event during their evolution (Jaillon et al., 2007; D’Hont et al., 2012; Tomato Genome Consortium, 2012). At a smaller scale, tandem and segmental duplications are also very common in angiosperms (Arabidopsis Genome Initiative, 2000; International Rice Genome Sequencing Project, 2005; Rizzon et al., 2006). Although the most common fate of duplicated genes is to be progressively lost, in some cases they can be retained in the genome, and adaptive as well as nonadaptive scenarios have been discussed to play a role in this preservation process (for review, see Moore and Purugganan, 2005; Hahn, 2009; Innan, 2009; Innan and Kondrashov, 2010). Whole-genome sequences also revealed that the same gene may undergo several rounds of duplication and retention. These LSE genes were shown to evolve under positive selection more frequently than single-copy genes in angiosperms (Fischer et al., 2014). That study analyzed general trends over whole genomes. Here, we ask if, and to what extent, this trend is observable at LRR-RLK genes. As this gene family is very dynamic and large, and in accordance with the results of Tang et al. (2010b), we expect the effect of positive selection to be even more pronounced than in the whole-genome average.We analyzed 33 Embryophyta genomes to investigate the evolutionary history of the LRR-RLK gene family in a phylogenetic framework. Twenty LRR-RLK SGs were identified, and from this data set, we deciphered the evolutionary dynamics of this family within angiosperms. The expansion/reduction rates were contrasted between SGs and species as well as in ancestral branches of the angiosperm phylogeny. We then focused on genes whose number increased dramatically in an SG- and/or species-specific manner (i.e. LSE genes). Those genes are likely to be involved in species-specific cellular processes or adaptive interactions and were used as a template to infer the potential occurrence of positive selection. This led to the identification of sites at which positive selection likely acted. We discuss our results in the light of angiosperm genome evolution and current knowledge of LRR-RLK functions. Positive selection footprints identified in LSE genes highlight the importance of combining evolutionary analysis and functional knowledge to guide further investigations.     

被子植物有性和无融合生殖过程中胼胝质壁的动态变化及其生物学意义

就被子植物有性生殖和无融合生殖过程中在某些特定时期和细胞中出现的胼胝质壁的活动情况作了介绍,并探讨了其生物学意义。