鹅观草居群表型多样性研究

应用居群生物学的原理和方法,对分布于中国7个地区、40个鹅观草(Roegneria kamojiOhwi)居群的表型多样性进行了分析。结果显示:鹅观草15个表型性状的总遗传多样性指数为1.825,表明鹅观草具有丰富的遗传多样性;居群内多样性指数为1.453,居群间为0.372,表型分化系数为28.38%,表明居群内的遗传变异是引起鹅观草变异的主要因素;在7个不同分布区,鹅观草的多样性存在较明显的差异,多样性指数从大到小的顺序为四川(1.583)>云南(1.392)>新疆(1.372)>甘肃(1.351)>内蒙古(1.284)>宁夏(1.235)>山西(0.890)。因此在进行鹅观草种质资源收集时,建议对四川和云南地区的鹅观草种质资源应给予较大关注。     

种间杂种染色体配对所揭示的披碱草属植物StY基因组分化及其进化意义

在同倍体(homoploid)植物杂交-分化的物种形成(speciation)过程中,杂交后代与亲本之间的有效生殖隔离是新物种形成的关键。杂种与亲本在时间、空间、生态环境和基因水平上的隔离,保证了杂种后代的分化和稳定,并逐渐形成新物种。为了研究披碱草属(Elymus)含StY基因组四倍体物种的系统演化关系,本文对来自亚洲不同地理分布区的26种披碱草属植物进行了大规模的种间杂交和杂种F1减数分裂染色体配对行为的分析。结果表明各物种之间有不同程度的杂交亲合力,杂交结实率在各杂交组合之间有较大的变异(在4．8％-100％之间);但各物种之间的杂种F1完全不育。证明各物种之间形成了明显的生殖隔离。种间杂种F1减数分裂中期-I染色体配对分析的结果进一步表明,StY基因组随各披碱草属物种地理分布的不同而有不同程度的分化。来自同一分布区(如东亚或西亚分布区之内)物种的StY基因组分化程度较低,但来自不同分布区(如东亚和西亚)物种之间相同的StY基因组具有显著的分化。表明地理隔离对含StY基因组物种的分化起到了十分重要的作用。通过对种间和种内杂种F1的减数分裂异常现象和染色体配对频率变化规律的分析,作者认为细胞学水平的变化,如基因组同源性的分化和染色体结构的变异等都在杂种后代与亲本之间产生生殖隔离并逐渐形成新物种的进化过程中起到了积极的作用。     

Iris japonica×Iris confusa种间杂种的细胞遗传学研究

对Irisjaponica、I confusa和它们的人工种间杂种花粉母细胞减数分裂中期Ⅰ染色体配对行为、形态学和繁育特征进行了分析 ,结果表明 :(1)I japonica和I confusa的形态差异较小 ,杂种F1形态介于亲本之间 ;(2 )人工杂交比较容易 ,杂种种子能正常发育 ;杂种F1体细胞染色体数为 2n =30 ,减数分裂中期I染色体配对频率很高 ,为 13 9个二价体 ,构型为0 95Ⅰ + 5 95Ⅱ (棒形 ) + 7 95Ⅱ (环形 ) + 0 0 5Ⅲ + 0 2 7Ⅳ ,表明它们有相似的染色体组 ,亲缘关系很近 ;(3)杂种中有少量的三价体和四价体存在 ,可能是亲本染色体间发生了结构重排或是部分同源染色体配对 ;(4)杂种中大多数细胞存在 2个单价体 ,有的高达 8个单价体 ;花粉育性为 5 0 5 1% ,不能正常结实 ,表明I japonica和I confusa间存在生殖隔离 ,是独立的生物学物种 ;(5 )杂种F1减数分裂前期可观察到细胞融合 ,这可能是造成I japonica和I confusa不同居群出现多倍体和非整倍体的重要原因。     

鹅观草和簇毛麦属间杂种的形态学和细胞遗传学研究

通过幼胚离体拯救培养,成功地获得鹅观草Roegneria kamoji Ohwi(2n=6x=42,StStHHYY)和 簇毛麦Dasypyrum villosum(L．)Candargy(2n=2x=14,VV)属间杂种,对这两个种及其杂种F_l的形 态学、繁育学和减数分裂配对行为进行了研究。结果表明：(1)鹅观草和簇毛麦形态差异极大,杂种F1 的形态特征介于父母本之间;(2)杂交结实率低,为11．63％;经幼胚组织培养,获2株杂种苗;杂种F_l高 度不育,表明亲本间存在极强的生殖隔离,是独立的生物学种;(3)杂种F_l体细胞染色体数目为稳定的 28条,减数分裂中期I染色体配对很低,其构型为：26．72Ⅰ+0．62Ⅱ+0．02Ⅲ。表明鹅观草的St、H、Y染色体组与簇毛麦的V染色体组部分同源性很低,它们的亲缘关系较远。     

垂珠花自然居群表型性状及遗传多样性分析

为探索垂珠花自然居群间表型差异的内在因素,该研究用14个表型性状和微卫星分子标记法对6个垂珠花自然居群进行遗传多样性分析。结果表明:(1)14个表型性状在居群间和居群内皆存在极显著差异;居群间平均表型分化系数(V_(st))为22.64%,垂珠花自然居群内的表型变异大于居群间的表型变异,说明居群内变异是垂珠花居群的主要变异来源。(2)4对微卫星引物在87个个体上共检测到43个等位基因,平均每条引物10.63个,平均多态位点百分率(P)为100%;平均观察杂合度(H_o)和平均期望杂合度(H_e)分别为0.659和0.811,Shannon多样性指数(I)平均达1.894;居群内遗传多样性(H_s)高达0.811,居群间遗传多样性(D_(st))仅为0.110,居群间遗传分化系数(F_(st))为0.118,说明11.8%的遗传变异来自于居群间,88.2%的遗传变异来自于居群内,与表型变异研究所得结论大体相一致。表型性状和微卫星分子标记结果均表明垂珠花自然居群存在丰富的遗传变异,较高的基因流(Nm=2.050)使得垂珠花居群间的遗传变异小于居群内的遗传变异。     

普通小麦与鹅观草属间杂种F1及BC1的分子细胞遗传学、育性和赤霉病抗性研究

通过胚拯救, 成功获得鹅观草Roegneria kamoji (2n=6x=42, SSHHYY)和普通小麦中国春Triticum aesti-vum (2n=6x=42, AABBDD)的正反交属间杂种F1, 并对这些杂种F1及其BC1的形态学、减数分裂配对行为、育性和赤霉病抗性进行研究。结果表明, (鹅观草×中国春)F1和(中国春×鹅观草)F1的形态介于双亲之间。杂种F1花粉母细胞减数分裂中期I染色体构型分别为40.33I + 0.78II + 0.03III和40.40I + 0.79II 。杂种F1高度雄性不育, 用中国春花粉与其回交可获得BC1代种子。(鹅观草×中国春) F1×中国春BC1植株的染色体数目主要分布在55~63之间, 单价体较多, 植株高度不育; (中国春×鹅观草)F1×中国春BC₁植株染色体数目也主要分布在55~63之间, 但其中部分植株拥有整套小麦染色体且能正常配对、分离, 可形成部分可育花粉粒, 能收到少量自交结实种子。在 (鹅观草×中国春)F1中有1株穗型趋向中国春, 其染色体数目为2n=63, 经染色体分子原位杂交(GISH)检测, 含有42条小麦染色体和21条鹅观草染色体。该杂种F1在减数分裂中期I平均每个花粉母细胞有26.40I+18.30II, 但植株高度雄性不育, 用中国春花粉回交能收到BC₁种子。(鹅观草×中国春) F₁ (2n=63)×中国春BC₁的染色体数目主要分布在40~59之间, 其中的外源染色体已经逐渐减少, 虽然该BC₁的穗型已接近中国春, 但仍然高度不育。赤霉病抗性鉴定结果显示, 所有杂种F1及大部分BC₁对赤霉病均表现出较好的抗性。     

余甘子天然居群果实形态变异研究

为了从数量性状上分析余甘子天然居群果实表型性状在居群间和居群内的变异,在云南省选取了6个余甘子天然居群,分别测量了其8个果实表型性状并作比较分析。结果显示:(1)余甘子果实表型性状在居群间和居群内存在着较丰富的变异,8个果实性状在居群间的F值为20.17～35.21,部分差异达到显著水平;居群内的F值为1.27～3.07,差异均达到极显著水平。(2)余甘子居群内的平均表型变异接近90%,是余甘子果实表型遗传变异的主要来源。(3)余甘子表型性状的变化与生态和地理因子显著相关;主成分分析结果显示,单果重、果实横径、外果皮重、果核宽、果核重等5个表型性状指标是反映滇西北余甘子果实表型差异的主要因素。     

普通小麦与鹅观草属间杂种F1及BC1的分子细胞遗传学、育性和赤霉病抗性研究

通过胚拯救, 成功获得鹅观草Roegneria kamoji (2n=6x=42, SSHHYY)和普通小麦中国春Triticum aesti-vum (2n=6x=42, AABBDD)的正反交属间杂种F1, 并对这些杂种F1及其BC1的形态学、减数分裂配对行为、育性和赤霉病抗性进行研究。结果表明, (鹅观草×中国春)F1和(中国春×鹅观草)F1的形态介于双亲之间。杂种F1花粉母细胞减数分裂中期I染色体构型分别为40.33I + 0.78II + 0.03III和40.40I + 0.79II 。杂种F1高度雄性不育, 用中国春花粉与其回交可获得BC1代种子。(鹅观草×中国春) F1×中国春BC1植株的染色体数目主要分布在55~63之间, 单价体较多, 植株高度不育; (中国春×鹅观草)F1×中国春BC₁植株染色体数目也主要分布在55~63之间, 但其中部分植株拥有整套小麦染色体且能正常配对、分离, 可形成部分可育花粉粒, 能收到少量自交结实种子。在 (鹅观草×中国春)F1中有1株穗型趋向中国春, 其染色体数目为2n=63, 经染色体分子原位杂交(GISH)检测, 含有42条小麦染色体和21条鹅观草染色体。该杂种F1在减数分裂中期I平均每个花粉母细胞有26.40I+18.30II, 但植株高度雄性不育, 用中国春花粉回交能收到BC₁种子。(鹅观草×中国春) F₁ (2n=63)×中国春BC₁的染色体数目主要分布在40~59之间, 其中的外源染色体已经逐渐减少, 虽然该BC₁的穗型已接近中国春, 但仍然高度不育。赤霉病抗性鉴定结果显示, 所有杂种F1及大部分BC₁对赤霉病均表现出较好的抗性。     

Y基因组分化及其进化意义

在同倍体（homoploid）植物杂交－分化的物种形成（speciation）过程中,杂交后代与亲本之间的有效生殖隔离是新物种形成的关键。杂种与亲本在时间、空间、生态环境和基因水平上的隔离,保证了杂种后代的分化和稳定,并逐渐形成新物种。为了研究披碱草属（Elymus）含S_tY基因组四倍体物种的系统演化关系,本文对来自亚洲不同地理分布区的26种披碱草属植物进行了大规模的种间杂交和杂种F₁减数分裂染色体配对行为的分析。结果表明各物种之间有不同程度的杂交亲合力,杂交结实率在各杂交组合之间有较大的变异（在4.8%-100%之间）;但各物种之间的杂种F₁完全不育。证明各物种之间形成了明显的生殖隔离。种间杂种F₁减数分裂中期-I染色体配对分析的结果进一步表明,S_tY基因组随各披碱草属物种地理分布的不同而有不同程度的分化。来自同一分布区（如东亚或西亚分布区之内）物种的S_tY基因组分化程度较低,但来自不同分布区（如东亚和西亚）物种之间相同的S_tY基因组具有显著的分化。表明地理隔离对含S_tY基因组物种的分化起到了十分重要的作用。通过对种间和种内杂种F₁的减数分裂异常现象和染色体配对频率变化规律的分析,作者认为细胞学水平的变化,如基因组同源性的分化和染色体结构的变异等都在杂种后代与亲本之间产生生殖隔离并逐渐形成新物种的进化过程中起到了积极的作用。     

云南稻核心种质BC1F2代形态性状的遗传变异

在昆明孕穗期平均温度18.1℃的自然低温下,以云南当家品种滇屯502和合系35作轮回亲本,以548份云南稻核心种质为供体亲本配制的杂种BC1F2(458份)为材料,按籼粳、稻作区进行云南稻核心种质杂种后代形态性状遗传变异研究。结果表明:(1)杂种BC1F2具有丰富的形态多样性,结实率、实粒数、秕粒数、穗颈长等形态性状都表现出较大的变异,平均多样性指数也较高。(2)籼粳间、稻作区间单一性状变异趋势基本一致,但多样性指数存在一定差异。     

Morphology,fertility and cytogenetics of intergeneric hybrid between Roegneria kamoji Ohwi and Dasypyrum villosum (L.) Candargy (Poaceae: Triticeae)

The intergeneric hybrids between Roegneria kamoji Ohwi and Dasypyrum villosum (L.)Candargy were successfully obtained by means of embryo culture in vitro. Studies on morphology, fertility and chromosome pairing behavior in meiosis of the parents and their hybrid F_l were carried out in the present work. The results showed that: (1) there were ob vious morphological differences between R. kamoji and D. villosum, and spikes of F_l plants were morphologically intermediate between the two parental species; (2) the seed set of the cross was 11.63%; the hybrid plant was infertile, which indicated that strong repro ductive isolation existed between the parents and R. kamoji and D. villosum were inde pendent biological species; (3) The somatic chromosome number in root-tips of F_l hybrids was 28. Chromosome pairing at MI of PMCs in F_l hybrids was quite low. The meiotic con figuration was 26.72 Ⅰ + 0.62 Ⅱ + 0.02 Ⅲ, which indicated that very low homoeology was detected between the St, H, Y genomes of R. kamoji and the V genome of D. villo- sum, and the relationship between the parental species was remote.     

Fitness of reciprocal F1 hybrids between Rhinanthus minor and Rhinanthus major under controlled conditions and in the field

The performance of first‐generation hybrids determines to a large extent the long‐term outcome of hybridization in natural populations. F₁ hybrids can facilitate further gene flow between the two parental species, especially in animal‐pollinated flowering plants. We studied the performance of reciprocal F₁ hybrids between Rhinanthus minor and R. major, two hemiparasitic, annual, self‐compatible plant species, from seed germination to seed production under controlled conditions and in the field. We sowed seeds with known ancestry outdoors before winter and followed the complete life cycle until plant death in July the following season. Germination under laboratory conditions was much lower for the F₁ hybrid formed on R. major compared with the reciprocal hybrid formed on R. minor, and this confirmed previous results from similar experiments. However, this difference was not found under field conditions, which seems to indicate that the experimental conditions used for germination in the laboratory are not representative for the germination behaviour of the hybrids under more natural conditions. The earlier interpretation that F₁ hybrid seeds formed on R. major face intrinsic genetic incompatibilities therefore appears to be incorrect. Both F₁ hybrids performed at least as well as and sometimes better than R. minor, which had a higher fitness than R. major in one of the two years in the greenhouse and in the field transplant experiment. The high fitness of the F₁ hybrids confirms findings from naturally mixed populations, where F₁ hybrids appear in the first year after the two species meet, which leads to extensive advanced‐hybrid formation and introgression in subsequent generations.     

Genetic analysis of hybrid seed formation ability of Brassica rapa in intergeneric crossings with Raphanus sativus

A hybridization barrier leads to the inability of seed formation after intergeneric crossings between Brassica rapa and Raphanus sativus. Most B. rapa lines cannot set intergeneric hybrid seeds because of embryo breakdown, but a B. rapa line obtained from turnip cultivar ‘Shogoin-kabu’ is able to produce a large number of hybrid seeds as a maternal parent by crossings with R. sativus. In ‘Shogoin-kabu’ crossed with R. sativus, developments of embryos and endosperms were slower than those in intraspecific crossings, but some of them grew to mature seeds without embryo breakdown. Intergeneric hybrid seeds were obtained in a ‘Shogoin-kabu’ line at a rate of 0.13 per pollinated flower, while no hybrid seeds were obtained in a line developed from Chinese cabbage cultivar ‘Chiifu’. F₁ hybrid plants between the lines of ‘Shogoin-kabu’ and ‘Chiifu’ set a larger number of hybrid seeds per flower, 0.68, than both the parental lines. Quantitative trait loci (QTLs) for hybrid seed formation were analyzed after intergeneric crossings using two different F₂ populations derived from the F₁ hybrids, and three QTLs with significant logarithm of odds scores were detected. Among them, two QTLs, i.e., one in linkage group A10 and the other in linkage group A01, were detected in both the F₂ populations. These two QTLs had contrary effects on the number of hybrid seeds. Epistatic interaction between these two QTLs was revealed. Possible candidate genes controlling hybrid seed formation ability in QTL regions were inferred using the published B. rapa genome sequences.     

鹅观草与大鹅观草杂种的细胞遗传学及形态学研究

大鹅观草（ Ｒｏｅｇｎｅｒｉａ ｇｒａｎｄｉｓ Ｋｅｎｇ） 是分布于我国陕西的一种多年生四倍体植物。为了探索大鹅观草与鹅观草（ Ｒｏｅｇｎｅｒｉａ ｋａｍｏｊｉ Ｏｈｗｉ） 间的物种生物学关系, 通过对其进行远缘杂交,幼胚离体培养, 合成了远缘杂种; 并对亲本及杂种Ｆ１ 代花粉母细胞减数分裂中期Ⅰ染色体配对行为及形态学进行了研究。结果表明杂种Ｆ１ 减数分裂染色体配对平均构型为： Ｒ－ ｋａｍｏｊｉ ×Ｒ－ｇｒａｎｄｉｓ１０－２６ Ⅰ＋ １２－３７ Ⅱ（ｃ－ 值＝ ０－７４） , 大鹅观草与鹅观草所含的ＳＹ 染色体组间存在较大的同源性分化; 杂种穗部特征大多数介于双亲之间。     

Increased pre-dispersal seed predation in sunflower crop-wild hybrids

The fitness of crop-wild hybrids can influence gene flow between crop and wild populations. Seed predation levels in crop-wild hybrid plants can be an important factor in determining plant fitness, especially in large-seeded crops such as sunflower. To determine patterns of pre-dispersal seed predation, seeds were collected from wild sunflowers (Helianthus annuus L.) and wild×crop F₁ hybrids at three experimental field sites in eastern Kansas. Seed heads were dissected and each seed was counted and scored for categories of seed damage by lepidopteran and coleopteran larvae. Hybrid seed heads showed significantly higher levels of insect-damaged seeds. The average hybrid plant had 36.5% of its seeds (or 45.1 seeds per plant) eaten by insect larvae while the average wild plant lost only 1.8% (or 95 seeds) to seed predators. Hybrid populations had higher levels of total insect damage even when date of flowering, flower head diameter, and the number of open heads within the study site were accounted for. These results suggest that the reduced fecundity of F₁ crop-wild sunflower hybrids demonstrated in other studies may be augmented by the increased seed predation in hybrid flower heads. Fecundity estimates of crop-wild hybrid and wild plants that disregard differential seed predation levels may not accurately reflect the actual relative contributions of hybrid and wild plants to future generations. Received: 21 December 1998 / Accepted: 8 July 1999     

分布于日本和中国的鹅观草及其杂种的形态学和细胞学研究

本文对分布于日本的Agropyron tsukushiense (Honda) Ohwi var. transiense (Hack.) Ohwi (2n=6x=42)和分布于中国的Roegneria kamoji Ohwi (2n=6x=42)及其杂种F_1(2n=6x=42)进行了形态学及细胞学的研究,并同时探讨了亲本种的亲缘关系。总体来看,亲本材料之间在形态上虽有差异但并不十分显著。杂种F_1的形态特征介于其父、母本之间。在减数分裂过程中,亲本种和杂种F_1的染色体配对行为均十分正常。但在检查了大量的成熟花粉和穗状花序之后,发现杂种F_1有部份不育现象。上述研究结果表明A. tsukushiense var. transiens的三个染色体组与R. kamoji的三个染色体组同源。结合形态学和育性等方面的研究资料,作者认为上述两个材料仍应属于同一分类等级。但必须指出,由于长期的地理隔离,他们之间产生了一定的形态变异和生殖障碍。按照国际植物命名法规(ICBN)上述两个材料应组合为:Roegneria tsukushiensis (Honda) B. R. Lu, Yen et J. L. Yang及其变种var. transiens (Hack.) B. R. Lu, Yen et J. L. Yang comb. nov.     

Geographical variation in postzygotic isolation and its genetic basis within and between two Mimulus species

The aim of this study is to investigate the evolution of intrinsic postzygotic isolation within and between populations of Mimulus guttatus and Mimulus nasutus. We made 17 intraspecific and interspecific crosses, across a wide geographical scale. We examined the seed germination success and pollen fertility of reciprocal F₁ and F₂ hybrids and their pure-species parents, and used biometrical genetic tests to distinguish among alternative models of inheritance. Hybrid seed inviability was sporadic in both interspecific and intraspecific crosses. For several crosses, Dobzhansky–Muller incompatibilities involving nuclear genes were implicated, while two interspecific crosses revealed evidence of cytonuclear interactions. Reduced hybrid pollen fertility was found to be greatly influenced by Dobzhansky–Muller incompatibilities in five out of six intraspecific crosses and nine out of 11 interspecific crosses. Cytonuclear incompatibilities reduced hybrid fitness in only one intraspecific and one interspecific cross. This study suggests that intrinsic postzygotic isolation is common in hybrids between these Mimulus species, yet the particular hybrid incompatibilities responsible for effecting this isolation differ among the populations tested. Hence, we conclude that they evolve and spread only at the local scale.     

Hybridization in Annual Plants: Patterns and DynamicsDuring a Four-Year Study in Mixed Rhinanthus Populations

Hybridization in annual plants is rare, but their short life cycle provides an excellent opportunity to study the dynamics of hybridization. Hybridization occurs between the annual hemiparasites Rhinanthus minor and Rhinanthus angustifolius (Orobanchaceae). Using flower morphology, Kwak (1980) found a prevalence of hybrids close to R. angustifolius in a single population. We aim to find whether this pattern is also found using genetic markers, whether it is generally occurring in mixed populations, and whether these populations are stable over time. We used species-specific genetic markers to determine the number of individuals in a range of hybrid classes in three mixed populations of different ages during four consecutive years. In the young population, F₁ hybrids were found in the first year and mostly hybrids between R. minor and these F₁s in the second year, but in the years after that, hybrids close to R. angustifolius became more abundant. We also found this in the two older populations, where hybrids close to R. angustifolius always occurred in higher frequencies than hybrids close to R. minor. Over time, R. angustifolius strongly increased in frequency in two populations. Patterns of marker presence and absence suggested that advanced-generation hybrids are mainly formed by backcrossing with one of the parents, predominantly R. angustifolius whenever its frequency in the population is higher than 15%. The dynamics of mixed populations depend on the ecological conditions that regulate the presence of the two parental species, and introgression into R. angustifolius seems prevalent.     

Genetic distances revealed by morphological characters,isozymes, proteins and RAPD markers and their relationships with hybrid performance in oilseed rape (<Emphasis Type="Italic">Brassica napus</Emphasis> L.)

Genetic distances (GDs) based on morphological characters, isozymes and storage proteins, and random amplified polymorphic DNAs (RAPD) were used to predict the performance and heterosis of crosses in oilseed rape (Brassica napus L.). Six male-sterile lines carrying the widely used Shaan2A cytoplasm were crossed with five restorer lines to produce 30 F₁ hybrids. These 30 hybrids and their parents were evaluated for seven agronomically important traits and their mid-parent heterosis (MPH) at Yangling, Shaanxi province in Northwest China for 2 years. Genetic similarity among the parents based on 34 isozyme and seven protein markers was higher than that based on 136 RAPDs and/or 48 morphological markers. No significant correlation was detected among these three sets of data. Associations between the different estimates of GDs and F₁ performance for some agronomic traits were significant, but not for seed yield. In order to enhance the predicting efficiency, we selected 114 significant markers and 43 favoring markers following statistical comparison of the mean values of the yield components between the heterozygous group (where the marker is present only in one parent of each hybrid) and the homozygous group (where the marker is either present or absent in both parents of each hybrid) of the 30 hybrids. Parental GD based on total polymorphic markers (GD_total, indicating general heterozygosity), significant markers (GD_sign, indicating specific heterozygosity) and favoring markers (GD_favor, indicating favoring-marker heterozygosity) were calculated. The correlation between GD_favor or GD_sign and hybrid performance was higher than the correlation between GD_total and hybrid performance. GD_sign and GD_favor significantly correlated with plant height, seeds per silique and seed yield, but not with the MPH of the other six agronomic traits with the exception of plant height. The information obtained in this study on the genetic diversity of the parental lines does not appear to be reliable for predicting F₁ yield and heterosis.