作物驯化的遗传学研究及其在大豆育种中的应用

驯化遗传学是进化生物学研究的一个重要分支,对作物驯化历程进行深入探索,可以更好地认识人类早期农耕文明的产生和发展,了解物种的形成乃至生命的起源和进化,对指导人们利用野生资源来改良现有的栽培作物和培育新的有价值的品种也具有重要的意义。本文综述了作物驯化的遗传学研究内容和进展,以及重要作物大豆驯化起源的研究概况,探讨了目前作物驯化研究对大豆驯化遗传学及野生大豆资源综合利用的指导意义。     

家养动物驯化起源的研究方法与进展

驯化动物伴随了人类上万年的进化历程。在驯化过程中, 家养动物表型和行为特征发生巨大改变, 并与人类生产生活的进步相适应。研究动物驯化问题对于了解遗传多样性及适应性进化, 解析复杂性状的遗传机制等具有重要的意义, 成为生物学领域的重点研究内容之一。本文专注于动物驯化的初始阶段, 首先介绍了驯化起源的时间与地点、驯化途径和驱动因、驯化后的扩散和品种选育; 其次从驯化对象的角度着重介绍了考古学、分子和群体遗传学两方面的家养动物驯化起源的研究策略、优势与不足, 以及未来的发展方向; 最后, 我们综合多方面的证据, 介绍了在中国驯化的家猪(Sus domesticus)和家鸡(Gallus gallus domesticus)以及其他主要家养动物考古和分子水平上驯化起源的研究进展。本文整合多种证据为家养动物驯化起源的研究提供了相对完整的视角和新的思路。     

水稻驯化与长江文明

水稻(即亚洲栽培稻Oryza sativa)是世界上最重要的粮食作物之一, 全球有超过半数以上人口以稻米为食。关于水稻是何时、何地、在什么环境下开始驯化等问题一直是学术界关注的热点。得益于分析技术的进步, 近年来考古学和遗传学研究在水稻驯化起源问题上取得了重要进展。本文简要综述了有关长江流域的水稻驯化起源的遗传学和考古学的研究进展, 并讨论了水稻驯化与稻作文化及长江文明的关系。遗传学研究结果认为水稻(粳稻)最早起源于中国长江流域及以南地区(珠江流域), 考古学证据则表明水稻最先于10,000-8,000 BP在中国长江流域被驯化, 水稻驯化和稻作农业的发展催生了长江文明。这些进展促进了我们对水稻驯化、稻作文化和长江文明的认识, 对长江流域重要植物资源的保护也有启示意义。     

家养动物的起源与驯化研究进展

动物的驯化是新石器时代农业革命的主要内容之一, 导致了人类生活方式从狩猎向畜牧的转变。开展家养动物起源与驯化的研究, 不仅有助于了解人类社会的发展史和人工选择下动物的进化过程, 而且有助于开展家养动物的良种选育和遗传多样性保护, 因此该研究领域受到了人们的长期关注。近年来, 随着更多考古证据的发现与分子遗传学的发展, 人们对于家养动物的起源与驯化有了更深入的认识。本文对家养动物起源与驯化研究中一些基本问题的研究方法和进展进行了综述。文章第一部分介绍了家养动物的野生祖先、起源地、起源时间、建群者大小和扩散路线等问题; 第二部分则涉及家养动物驯化的条件、过程、性状改变及其遗传机制等方面。最后指出了家养动物起源与驯化研究中依然存在的问题, 并对未来研究发展的趋势进行了讨论。     

家马的驯化起源与遗传演化特征

人类文明发展历史中, 家马(Equus ferus caballus)曾是推动文化交流、促进人类社会发展的主要动力。关于家马何时、何地被驯化以及在此过程中其遗传演化如何被人类影响等一直备受关注。近年来随着遗传学技术的发展, 人们对该问题有了更为深入的理解。本文回顾了近二十年来相关研究所取得的成果, 探讨了家马的驯化起源中心和驯化过程中的遗传演化特征, 并对未来的研究方向以及遗传资源保护提出了建议。分子标记遗传学和考古学研究认为家马可能来自多个驯化起源地种群, 然而最近的古DNA研究结果表明, 现代家马的驯化起源可能比之前人们所猜测的更加复杂, 古代博泰马被认为是最早被驯化的家马, 然而最近被证实并不是现代家马的直系祖先。如此复杂的驯化问题可能从多学科的层次才能解析清楚。人类社会活动直接或间接影响了家马的演化历程, 特别是工业革命以来家马的遗传基础发生了巨大变化, 其遗传多样性开始急剧衰退, 不少地方品种正逐渐走向衰落甚至灭绝。为确保农业生态安全不受威胁, 建议加强家马遗传资源保护与动物遗传学和文化地理之间的联系研究。     

主要农作物驯化研究进展与展望

农作物的驯化过程与农耕文明的发端密切相关,是作物种质资源遗传基础不断丰富衍化,利用价值不断完善的主要途径之一。本文重点从作物驯化的主要性状、基因组遗传区段、重要驯化基因的克隆、作物驯化后的传播与基因渗透、作物驯化理论与研究方法以及作物驯化研究的新趋势等方面评述近年来取得的主要研究进展,讨论研究存在的主要问题并对深入开展作物驯化研究进行展望。     

“银额果蝇及其B染色体研究”获云南省2000年科学技术二等奖

中国科学院昆明动物研究所凌发瑶等 ,在国家自然科学基金、中国科学院院长基金、日本文部省国际研究基金的支持下 ,从群体细胞遗传学和分子遗传学着手 ,对银额果蝇及其Ｂ染色体的起源、进化以及生物学效应等进行了系统深入的研究。发现我国银额果蝇自然群体内存在极其丰富的Ｂ染色体遗传资源 ,并认为这是一种十分难得的可供持续研究Ｂ染色体的最佳动物模型 ;同时还发现了银额果蝇自然群体进化过程中的中间过渡性新核型 ,这一发现对研究果蝇的系统进化以及染色体的进化方式有着重要意义。通过本项研究 ,查明了银额果蝇自然群体内存在Ｂ染色体…     

蚕豆种质资源、抗病育种和QTL定位及抗逆性研究进展

蚕豆是世界温带和亚热带地区一种重要的食用豆类作物,在中国的栽培历史超过2100年。中国是世界上蚕豆栽培面积最大、总产量最多的国家,蚕豆因其高效生物固氮、土壤改良和环境友好特性,已成为中国现代农业种植结构调整、西部经济欠发达地区和丘陵山区农民脱贫致富的重要经济作物。目前,多种DNA标记已广泛应用于大豆、菜豆、豌豆等豆类作物,并取得了一系列重要进展,但蚕豆分子遗传学的研究进展相对缓慢。本文对蚕豆的起源、分类、国内外蚕豆遗传多样性、遗传图谱构建,以及生长习性、抗病育种和QTL定位、抗逆性研究进行了综述,旨在为国内外蚕豆资源的深入研究和利用提供参考。     

黍稷的名实考证及规范

黍稷是起源于中国最古老的作物,糯者为黍,粳者为稷。长期以来对黍稷的称谓一直不能统一,特别是对稷的称谓在农史界一直争论不休,难以定论。本文从黍稷的起源演化过程、细胞学和遗传学的研究、黍稷是人类最早驯化的作物、我国甲骨文和古农书的记载等方面,论证了黍稷的起源早于粟。并以此为据,进一步辩证了稷不是粟,同时指出将黍稷称为软糜和硬糜、糜黍、糜子、黍子等不同称谓的不足之处,进而认为以黍稷作为规范称谓为好。     

江苏省植物细胞遗传学研究回顾与展望

20世纪初"遗传的染色体学说"的提出和证明标志着细胞遗传学交叉学科建立,伴随相关学科的发展,20世纪60年代末期细胞遗传学又与分子遗传学相结合,建立发展了分子细胞遗传学交叉学科。分子细胞遗传学以DNA分子原位杂交技术为核心,不断拓展应用领域,为生命科学研究提供了直观、高效的技术手段。原位杂交技术与基因组、细胞生物学等技术结合,被广泛应用于人类、动物、植物的起源、进化、驯化等基础研究和远缘杂交、染色体工程等应用研究。通过形象地展示DNA、RNA、蛋白质在细胞中的实际位置,揭示DNA序列之间的实际位置和顺序、亲缘物种间的进化关系和结构重排、基因组拼接序列的质量、转录水平RNA和翻译水平蛋白质的位置和数量变化等。江苏省遗传学会会员单位南京农业大学、扬州大学、南京林业大学、江苏师范大学、徐淮地区农科院等自20世纪中期开展细胞遗传学理论技术研究,伴随学科发展不断创新,建立了较完善的分子细胞遗传技术体系,并成功应用于开展植物系统进化、远缘杂交、染色体工程、基因组学等研究,取得了一批研究成果。本文将主要综述江苏省在该领域取得的重要进展,并展望未来发展方向。     

A single origin and moderate bottleneck during domestication of soybean (Glycine max): implications from microsatellites and nucleotide sequences

Background and Aims

It is essential to illuminate the evolutionary history of crop domestication in order to understand further the origin and development of modern cultivation and agronomy; however, despite being one of the most important crops, the domestication origin and bottleneck of soybean (Glycine max) are poorly understood. In the present study, microsatellites and nucleotide sequences were employed to elucidate the domestication genetics of soybean.

Methods

The genomes of 79 landrace soybeans (endemic cultivated soybeans) and 231 wild soybeans (G. soja) that represented the species-wide distribution of wild soybean in East Asia were scanned with 56 microsatellites to identify the genetic structure and domestication origin of soybean. To understand better the domestication bottleneck, four nucleotide sequences were selected to simulate the domestication bottleneck.

Key Results

Model-based analysis revealed that most of the landrace genotypes were assigned to the inferred wild soybean cluster of south China, South Korea and Japan. Phylogeny for wild and landrace soybeans showed that all landrace soybeans formed a single cluster supporting a monophyletic origin of all the cultivars. The populations of the nearest branches which were basal to the cultivar lineage were wild soybeans from south China. The coalescent simulation detected a bottleneck severity of K′ = 2 during soybean domestication, which could be explained by a foundation population of 6000 individuals if domestication duration lasted 3000 years.

Conclusions

As a result of integrating geographic distribution with microsatellite genotype assignment and phylogeny between landrace and wild soybeans, a single origin of soybean in south China is proposed. The coalescent simulation revealed a moderate genetic bottleneck with an effective wild soybean population used for domestication estimated to be ≈2 % of the total number of ancestral wild soybeans. Wild soybeans in Asia, especially in south China contain tremendous genetic resources for cultivar improvement.     

From crop domestication to super-domestication

Research related to crop domestication has been transformed by technologies and discoveries in the genome sciences as well as information-related sciences that are providing new tools for bioinformatics and systems' biology. Rapid progress in archaeobotany and ethnobotany are also contributing new knowledge to understanding crop domestication. This sense of rapid progress is encapsulated in this Special Issue, which contains 18 papers by scientists in botanical, crop sciences and related disciplines on the topic of crop domestication. One paper focuses on current themes in the genetics of crop domestication across crops, whereas other papers have a crop or geographic focus. One feature of progress in the sciences related to crop domestication is the availability of well-characterized germplasm resources in the global network of genetic resources centres (genebanks). Germplasm in genebanks is providing research materials for understanding domestication as well as for plant breeding. In this review, we highlight current genetic themes related to crop domestication. Impressive progress in this field in recent years is transforming plant breeding into crop engineering to meet the human need for increased crop yield with the minimum environmental impact - we consider this to be 'super-domestication'. While the time scale of domestication of 10 000 years or less is a very short evolutionary time span, the details emerging of what has happened and what is happening provide a window to see where domestication might - and can - advance in the future.     

Genome-wide signatures of the geographic expansion and breeding of soybean

Soybean is a leguminous crop that provides oil and protein. Exploring the genomic signatures of soybean evolution is crucial for breeding varieties with improved adaptability to environmental extremes. We analyzed the genome sequences of 2,214 soybeans and proposed a soybean evolutionary route, i.e., the expansion of annual wild soybean (Glycine soja Sieb. & Zucc.) from southern China and its domestication in central China, followed by the expansion and local breeding selection of its landraces (G. max (L.) Merr.). We observed that the genetic introgression in soybean landraces was mostly derived from sympatric rather than allopatric wild populations during the geographic expansion. Soybean expansion and breeding were accompanied by the positive selection of flowering time genes, including GmSPA3c. Our study sheds light on the evolutionary history of soybean and provides valuable genetic resources for its future breeding.

     

Population Structure and Domestication Revealed by High-Depth Resequencing of Korean Cultivated and Wild Soybean Genomes

Despite the importance of soybean as a major crop, genome-wide variation and evolution of cultivated soybeans are largely unknown. Here, we catalogued genome variation in an annual soybean population by high-depth resequencing of 10 cultivated and 6 wild accessions and obtained 3.87 million high-quality single-nucleotide polymorphisms (SNPs) after excluding the sites with missing data in any accession. Nuclear genome phylogeny supported a single origin for the cultivated soybeans. We identified 10-fold longer linkage disequilibrium (LD) in the wild soybean relative to wild maize and rice. Despite the small population size, the long LD and large SNP data allowed us to identify 206 candidate domestication regions with significantly lower diversity in the cultivated, but not in the wild, soybeans. Some of the genes in these candidate regions were associated with soybean homologues of canonical domestication genes. However, several examples, which are likely specific to soybean or eudicot crop plants, were also observed. Consequently, the variation data identified in this study should be valuable for breeding and for identifying agronomically important genes in soybeans. However, the long LD of wild soybeans may hinder pinpointing causal gene(s) in the candidate regions.     

中国野生大豆(Glycine soja)遗传资源主要形态、遗传变异和结构

介绍了我国野生大豆遗传资源主要的形态类型、物种内遗传关系和遗传结构。进化的瓶颈不仅发生在由野生大豆到栽培大豆,也以另一种方式"分化瓶颈"出现于同性状的不同表型类型之间。野生大豆种内种子大小类型是否存在遗传分化?野生和半野生大豆的边界在哪?半野生大豆如何产生的?半野生大豆遗传上密切于栽培种还是野生种?百粒重3～4g的小粒半野生大豆与百粒重8.5g以上的特大粒半野生大豆是否有遗传差异?百粒重8.5g以上的特大粒半野生大豆是否属于栽培大豆?野生大豆的种皮色和种子大小哪个更能反映进化程度?栽培大豆基因是否已经渗入到野生大豆?对这些在学术界常年存在的疑问本文介绍了我们的研究答案。我们认为"真"半野生大豆不存在于现在中国半野生资源收集品中;一些野生大豆中的白花、灰毛、无泥膜性状来源于栽培大豆的基因渗透。     

长江流域野生猕猴桃遗传资源的潜在价值、现状分析与保护策略

长江流域是猕猴桃属(Actinidia)植物起源和演化的关键分布区, 富集了全世界重要的猕猴桃属野生物种资源和中华-美味猕猴桃物种复合体(Actinidia chinensis-A. deliciosa species complex)的种群遗传资源。作为水果作物, 猕猴桃在100多年前从长江流域经上海到新西兰, 通过栽培、驯化, 逐渐发展成新兴果树产业。目前, 猕猴桃植物的研究已在二倍体“红阳”中华猕猴桃(Actinidia chinensis cv. Hongyang)的基因组测序、种间关系的重测序分析和基础的分子系统发育、种群遗传结构等方面取得长足进步, 但基于最新研究成果的基本资源评价还相当匮乏, 对猕猴桃野生资源的保护与可持续利用亟待加强。本文回顾了栽培猕猴桃的驯化简史与猕猴桃属植物系统分类的研究进展, 通过与其他流域的比较, 对长江流域野生猕猴桃资源的潜在价值和现状进行分析, 阐述了该流域猕猴桃属植物的分布特点和受威胁的状况, 并针对目前存在的问题提出建立长效的保护机制、加强遗传资源的基础科研调查和系统评价, 以及健全种质资源保存规范和促进可持续利用等相应保护的策略。     

Variation and selection at the CAULIFLOWER floral homeotic gene accompanying the evolution of domesticated Brassica oleracea

The evolution of plant morphologies during domestication events provides clues to the origin of crop species and the evolutionary genetics of structural diversification. The CAULIFLOWER gene, a floral regulatory locus, has been implicated in the cauliflower phenotype in both Arabidopsis thaliana and Brassica oleracea. Molecular population genetic analysis indicates that alleles carrying a nonsense mutation in exon 5 of the B. oleracea CAULIFLOWER (BoCAL) gene are segregating in both wild and domesticated B. oleracea subspecies. Alleles carrying this nonsense mutation are nearly fixed in B. oleracea ssp. botrytis (domestic cauliflower) and B. oleracea ssp. italica (broccoli), both of which show evolutionary modifications of inflorescence structures. Tests for selection indicate that the pattern of variation at this locus is consistent with positive selection at BoCAL in these two subspecies. This nonsense polymorphism, however, is also present in both B. oleracea ssp. acephala (kale) and B. oleracea ssp. oleracea (wild cabbage). These results indicate that specific alleles of BoCAL were selected by early farmers during the domestication of modified inflorescence structures in B. oleracea.     

Brassica rapa Domestication: Untangling Wild and Feral Forms and Convergence of Crop Morphotypes

The study of domestication contributes to our knowledge of evolution and crop genetic resources. Human selection has shaped wild Brassica rapa into diverse turnip, leafy, and oilseed crops. Despite its worldwide economic importance and potential as a model for understanding diversification under domestication, insights into the number of domestication events and initial crop(s) domesticated in B. rapa have been limited due to a lack of clarity about the wild or feral status of conspecific noncrop relatives. To address this gap and reconstruct the domestication history of B. rapa, we analyzed 68,468 genotyping-by-sequencing-derived single nucleotide polymorphisms for 416 samples in the largest diversity panel of domesticated and weedy B. rapa to date. To further understand the center of origin, we modeled the potential range of wild B. rapa during the mid-Holocene. Our analyses of genetic diversity across B. rapa morphotypes suggest that noncrop samples from the Caucasus, Siberia, and Italy may be truly wild, whereas those occurring in the Americas and much of Europe are feral. Clustering, tree-based analyses, and parameterized demographic inference further indicate that turnips were likely the first crop type domesticated, from which leafy types in East Asia and Europe were selected from distinct lineages. These findings clarify the domestication history and nature of wild crop genetic resources for B. rapa, which provides the first step toward investigating cases of possible parallel selection, the domestication and feralization syndrome, and novel germplasm for Brassica crop improvement.