浅析物种概念的演变历史

本文是一篇关于物种概念演变的简述。生物学家用不同的方法或标准划分物种, 就形成了不同的物种概念, 如生物学物种、形态学物种、生态学物种、进化物种、系统发生或支序物种, 或它们的组合, 等等。它们都揭示了物种属性的特定侧面, 都是不同物种客观存在的真实反映, 但都无法令所有人满意。对真核生物来说, 无论它们在形态上的差别有多大, 生殖隔离(不能产生可育的后代)应该是两个群体能否真正分化成不同物种的关键, 这种隔离机制可以是地理的、行为的或其他方式; 而生殖隔离总会伴随着一些形态或遗传上的变化, 虽然这些特征可能与生殖隔离本身并无多大关系, 但往往成为分类学家或分子进化生物学家区分种的依据,对已经灭绝的化石物种来说, 生殖隔离的物种划分方式就无能为力了。如何准确定义一个物种依然充满着矛盾, 因为基于生殖隔离的物种概念不实用, 而实用的物种概念(如形态学物种)又被认为是人为的。     

菌物分类学研究中常见的物种概念

物种是生物多样性与分类学研究的基本单元, 物种识别是生物学研究的基本问题之一。物种的划分一直以来都没有一个明确统一的标准, 这使得分类学多少带有主观的色彩, 并经常被看作艺术而不完全是科学的研究。本文简要概述了菌物分类学研究中常见的3个物种概念, 即形态学种、生物学种和系统发育学种的背景和应用现状, 并通过实例讨论了这3个物种概念的特点及应用中存在的问题, 特别是各个物种概念之间的交错, 以期为菌物分类学研究和物种概念探讨提供参考。     

系统遗传学与合成生物学——21世纪的生物工程产业化

基因型-表现型复杂生物系统由多基因群调控,细胞发生的信号传导路径、多基因相互作用与细胞系谱定位形成生物系统的结构-图式发生遗传学,但分子、细胞和器官的结构、图式形成机理还不很清楚。复杂生物系统的图式演化是细胞的物种进化、细胞形态发育的细胞发生非线性动力学过程,包括：1）物种基因组结构内等位基因替代构成物种内基因多样性调控;2）物种间进化的基因组结构层次级别的自组织化。系统理论应用于系统生态学（Van Dyne GM．1966）、系统生理学（Sagawa K．1973）、系统心理学（Titchener EB．1992）、系统生物医学（Kamada T．1992）、系统生物学（zieglgansherger W,Tolle TR．1993）、系统生物工程与系统遗传学（Zengg：BJ.1994）的建立,以及遗传学机理的生物系统分析。细胞的基因组结构自组织化形成生物的系统发生,基因组的结构变化形成物种的适应变异,生物体结构的基因组复制与表达的细胞自组织化构成生物个体发生。基于系统遗传学的工程应用,合成生物学探索生物系统泛进化,包括人工生物体的遗传工程、基因调控和仿生智能的纳米生物机器,构成生物系统的人工引导进化。     

“物种”与“个体”:究竟谁是生物多样性保护的恰当对象?

生物多性保护实践涉及到两大基本问题:一是我们要保护的是什么,二是我们优先保护什么。从保护对象的角度看,物种和个体均不是生物多样性保护的恰当对象,准确的目标是“物种多样性”。所有的物种都在物种分化程度与分化时间上千差万别,从而导致不可能有统一的标准去划分不同的物种。从多样性或者说物种多样性的目标出发,作为保护对象的物种必须是独立进化的单元,同时,它必须有一个可操作的、相对合理的界定方法,而整合分类法可能就是一条极具吸引力的道路。生物分类学家应当尽可能利用整合分类思想和整合方法,建立可操作的、相对合理的物种概念和优先保护级别划分体系,以避免实际工作中的冲突与困境,更好地服务于生物多样性保护事业。     

论物种的客观真实性

关于物种的概念和本质的争论由来已久。大多数学者认为种是自然界客观存在的真实单位,是形态上和生殖上间断的群体体系,与种内连续的群体体系——宗,有着本质的区别。少数学者则认为种是主观意识的产物,是为了分类的目的而任意确定的人为的分类阶元,与宗只有程度的不同而无本质的区别;只有个体才是客观真实的单位。以形态标准为主的分类学种概念和以生殖隔离为标准的生物学种概念只能说代表了自然界有机体类群不连续性的两个不尽相符,但部分一致的侧面,而不能说就是客观物种的反映。自然界的生命有机体有两重性。它们既不是一群乱七八糟的乌合之众,也不是秩然不紊的鳞次栉比,而是以群体的组织形式适应于各自分布区内的生态位,在表型和基因型的变异式样上具有一定规律的遗传多样性。物种的客观本质也有两重性。物种作为有机体组合的单位既存在又不存在。说物种客观存在是因为按形态或生殖的不连续性标准划分的“种”在自然界具有一定的普遍性;说物种并非客观存在是因为至今还没有一个能把所有的生命有机体都划分成统一的生物学单位的不连续性标准。     

物种与物种多样性

本文首先讨论生物物种的科学概念和生物学本质，分析物种客观存在的自然属性和物种概念的局限性，认为物种的生物学属性和物种多样性的科学属性之间有着本质联系。物种多样性研究的实质是研究生物物种的生物学多样性。度量物种多样性程度有多种方法，但物种数目是物种多样性程度最直接、也是最基本的表达，估计物种多样性数目是当前国际上物种多样性研究的核心与热点内容。物种多样性产生的根源是物种形成，物种绝灭速率是维持物种多样性的关键因素。本文简要总结了物种形成与绝灭的基本模式和机制，通过分析生物地理区系与物种多样性研究的密切关系，说明物种的区系成份分析是物种多样性大尺度格局研究的重要内容。     

进化发育生物学--发育、进化和遗传的再联合

发育生物学和进化生物学,以及遗传学历史上曾一度是彼此不分的统一体,后来由于各自研究重点的不同和相应研究手段的独立发展彼此分道扬镳了。如今,由于分子遗传学研究手段的革新使得基因序列测定成为分析发育机理、区分物种和评估种间亲缘关系的常规手段,三者又在基因水平上再度统一起来了,并形成一门被称为进化发育生物学（ｅｖｏｌｕｔｉｏｎａｒｙ ｄｅｖｅｌｏｐｍｅｎｔａｌ ｂｉｏｌｏｇｙ）的新学科。     

细胞水平的分类学——细胞分类学

各学科互相渗透形成众多的边缘学科,这是当代科学发展的重要特点。细胞学和细胞遗传学同分类学之间的渗透,出现了细胞水平的分类学——细胞分类学。形态-地理学标准是传统分类学的主要方法。本世纪以来,由于实验生物学的蓬勃发展,分类学曾一度遭到冷落。然而,实验对象必须正确鉴定,实验所得信息需要记录和储存在确切的信息库内;自然界又发现不少兄弟种(sibling species)它们在形态上不能或难以区别,但却有被保护的独立的基因库。因此形态-地理学的物种概念被生物学物种概念所替代。与此同时,实验生物学,特别是     

我国植物种级水平分类学研究刍议

对洪德元先生最近在《生物多样性》(2016年第24卷第3期)发表的《关于提高物种划分合理性的意见》一文中的部分观点进行了进一步阐述。强调我国植物确实还存在大量种级水平的分类学问题有待解决, 我国植物分类学研究在一些重要发展阶段(如系统阶段和物种生物学阶段)上存在明显缺失, 需要弥补。指出分类学发展到今天, 不宜再强调“经典分类学”和“实验分类学”之分, 应采用多学科手段解决分类学问题; 我国应加强植物分类专著水平的研究工作, 注意培养年轻一辈植物分类学专著工作者; 在分类处理中应用居群概念和统计学方法时应特别谨慎; 在系统植物学中接受物种概念的多元性是必要的, 但要向达到广义的生物学种概念努力, 不宜以有所谓的“归并派”和“细分派”之分为借口而完全主观地划分物种。     

路径依赖下的物种形成机制

生物科学几乎所有研究都需要物种概念作为基础, 生物多样性研究亦需要可操作的物种概念, 但现有物种概念存在不同程度的人为因素或难操作性, 对物种划分造成不利影响。本文引入“进化路径”这一概念, 说明适合度景观时刻变化着, 物种在每个进化时间点上依据瞬时适合度选择下一时刻的进化状态, 且总是沿着动态适合度景观中适合度增加的方向进化。基于演化博弈的方法, 以随机过程为例模拟物种的进化过程。进而提出路径依赖下的物种形成机制, 并在此基础上给出可操作的物种定义, 即: 针对基因、性状、生态过程等任一状态下两个群体内个体的多个变量做统计分析, 若群体之间同时在两个或多个维度状态下呈现出的不连续性d大于群体内变量呈现出的差异性σ_k, 则拥有相应变量的个体属于不同物种。     

The subspecies concept in butterflies: has its application in taxonomy and conservation biology outlived its usefulness?

Subspecies lie at the interface between systematics and population genetics, and represent a unit of biological organization in zoology that is widely used in the disciplines of taxonomy and conservation biology. In this review, we explore the utility of subspecies in relation to their application in systematics and biodiversity conservation, and briefly summarize species concepts and criteria for their diagnosis, particularly from an invertebrate perspective. The subspecies concept was originally conceived as a formal means of documenting geographical variation within species based on morphological characters; however, the utility of subspecies is hampered by inconsistencies by which they are defined conceptually, a lack of objective criteria or properties that serve to delimit their boundaries, and their frequent failure to reflect distinct evolutionary units according to population genetic structure. Moreover, the concept has been applied to populations largely comprising different components of genetic diversity reflecting contrasting evolutionary processes. We recommend that, under the general lineage (unified) species concept, the definition of subspecies be restricted to extant animal groups that comprise evolving populations representing partially isolated lineages of a species that are allopatric, phenotypically distinct, and have at least one fixed diagnosable character state, and that these character differences are (or are assumed to be) correlated with evolutionary independence according to population genetic structure. Phenotypic character types include colour pattern, morphology, and behaviour or ecology. Under these criteria, allopatric subspecies are a type of evolutionarily significant unit within species in that they show both neutral divergence through the effects of genetic drift and adaptive divergence under natural selection, and provide an historical context for identifying biodiversity units for conservation. Conservation of the adaptedness and adaptability of gene pools, however, may require additional approaches. Recent studies of Australian butterflies exemplify these points. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Why the Phylogenetic Species Concept?—Elementary

Although species play a number of unique and necessary roles in biology, none are more important than as the elements of phylogeny, nomenclature, and biodiversity study. Species are not divisible into any smaller units among which shared derived characters can be recognized with fidelity. Biodiversity inventory, assessment, and conservation are dependent upon a uniformly applicable species concept. Species are the fundamental units in formal Linnaean classification and zoological nomenclature. The Biological Species Concept, long given nominal support by most zoologists, forced an essentialy taxonomic problem (what are species?) into a population genetics framework (why are there species?). Early efforts at a phylogenetic species concept focused on correcting problems in the Biological Species Concept associated with ancestral populations, then applying phylogenetic logic to species themselves. Subsequently, Eldredge and Cracraft, and Nelson and Platnick, each proposed essentially identical and truly phylogenetic species concepts that permitted the rigorous recognition of species prior to and for the purposes of phylogenetic analysis, yet maintained the integrity of the Phylogenetic Species Concept outside of cladistic analysis. Such phylogenetic elements have many benefits, including giving to biology a unit species concept applicable across all kinds of living things including sexual and asexual forms. This is possible because the Phylogenetic Species Concept is based on patterns of character distributions and is therefore consistent with the full range of possible evolutionary processes that contribute to species formation, including both biotic and abiotic (even random) factors.     

Species Concept in Primates

The way we view the Species category in Primates, as in other animals, especially other vertebrates, has been going through a revolution over the past 20 years or so. Much is wrong with the idea that we can define species according to whether or not they are “reproductively isolated”: this concept, the so‐called Biological Species Concept, has never offered any guidelines in the case of allopatric populations; this has now been shown to be simply wrong. Although other ways of looking at species – the Evolutionary, Recognition, Cohesion and Genetic Species Concepts – have all provided particular insights, the only proposal to offer a repeatable, falsifiable definition of species is the Phylogenetic Species Concept. This has been criticised for increasing the number of species to be recognised, although it is not clear why this should be a problem: indeed, it tells us that the world is far richer in biodiversity than we had conceived. Am. J. Primatol. 74:687‐691, 2012. © 2012 Wiley Periodicals, Inc.     

Analysis of biodiversity across levels of biological organization: a problem of defining traits

Biodiversity is a term that comprises the appearance, structure and function of all levels of biological organization, including genes, species and ecosystems. The vast majority of measures of biodiversity (usually termed ‘diversity indices’) considers only number, proportion and distribution of species which belong to a specified group and exist in a defined area or ecosystem. Genetic diversity as a part of biodiversity within species (or populations) was either not regarded in this respect or was treated (by geneticists) as a separate entity of diversity quantified with separate measures. Little attention has been given to the integration of both types of diversity, within and among species, in a single measurement (termed ‘transspecific’ diversity). In order to attain this integration on a general basis, an operational trait concept is developed which allows the determination of variation in traits observable in members not only of the same species but also of different species. The concept rests on methods of investigation that can be adapted to a broader range of organisms without modification of their characteristics. Once a trait is specified on this basis, any meaningful measure of diversity can be applied to assess biodiversity across levels of biological organization. The utility of the concept is demonstrated by application to the results of an earlier study on associations between species and genetic diversity in a forest tree community. Attributes of isozymes which are visible in electrophoresis are used as a transspecific genetic trait.     

Detecting cryptic species in sympatry and allopatry: analysis of hidden diversity in Polyommatus (Agrodiaetus) butterflies (Lepidoptera: Lycaenidae)

Modern multilocus molecular techniques are a powerful tool in the detection and analysis of cryptic taxa. However, its shortcoming is that with allopatric populations it reveals phylogenetic lineages, not biological species. The increasing power of coalescent multilocus analysis leads to the situation in which nearly every geographically isolated or semi‐isolated population can be identified as a lineage and therefore raised to species rank. It leads to artificial taxonomic inflation and as a consequence creates an unnecessary burden on the conservation of biodiversity. To solve this problem, we suggest combining modern lineage delimitation techniques with the biological species concept. We discuss several explicit principles on how genetic markers can be used to detect cryptic entities that have properties of biological species (i.e. of actually or potentially reproductively isolated taxa). Using these principles we rearranged the taxonomy of the butterfly species close to Polyommatus (Agrodiaetus) ripartii. The subgenus Agrodiaetus is a model system in evolutionary research, but its taxonomy is poorly elaborated because, as a rule, most of its species are morphologically poorly differentiated. The taxon P. (A.) valiabadi has been supposed to be one of the few exceptions from this rule due to its accurately distinguishable wing pattern. We discovered that in fact traditionally recognized P. valiabadi is a triplet of cryptic species, strongly differentiated by their karyotypes and mitochondrial haplotypes.     

The What,Why and How of Primate Taxonomy

Taxonomy has a well-defined role, which is much more than simply stamp-collecting and pigeon-holing. Species are the units of classification, biogeography and conservation; as such they must be defined as objectively as possible. The biological species concept, still widely used in biology, though predominantly by non-taxonomists and all too often misunderstood, is a process-based concept, which offers no criterion for the classification of allopatric populations beyond inference and hypothesis. The phylogenetic species concept—a pattern-based concept—is as nearly objective as we are likely to get. Amount of difference is not a criterion for recognizing species. It is not possible to insist on monophyly at the specific level, but it is mandatory for the higher categories (genus, family, etc.). The rank we assign to a given supraspecific category should be determined by its time depth.     

基于生物学物种定义探讨物种形成理论与验证的研究进展

物种形成是进化生物学研究的一个永恒主题, 由于生物群体进化是连续和动态的, 物种界限变得难于界定。本文首先讨论了3种地理物种形成模式(同域、邻域及异域), 并分析了近期报道的研究证据。其次, 评述了合子后生殖隔离机制的分子遗传基础和应用群体基因组数据分析的证据, 包括BDMI模型(Bateson-Dobzhansky-Muller incompatibility)、QTLs (quantitative trait loci)、霍尔丹法则及大X染色体效应。最后, 探讨了交配系统作为合子前隔离机制之一与物种形成的关系, 认为近交或自交通过扩大种群遗传结构分化, 增强不同交配系统的种群间不对称基因渐渗, 或种群间无基因渐渗等途径, 促进新物种形成。已知植物交配系统的演化更倾向于从异交(或自交不亲和)向自交(或近交亲和)方式, 花性状和基因组的分化推动形成所谓的自交综合征, 研究交配系统驱动或强化物种形成模式对认识植物物种形成机制有重要意义。