以礼草属的分类研究

以礼草属Kengyilia Yen et J. L. Yang 是禾本科Poaceae小麦族trib．Triticeae中新近建立的屑,针对其存在问题,对它进行了全面清理,提出了一个新的分类系统。新系统含3组、26种、6变种,其中3个组为新设立,并包括1新种、7个新等级、新组合及新异名。此外,还报道了一些类群的分布新记录。     

A taxonomical study on the genus Kengyilia Yen et J. L. Yang

Kengyilia Yen et J L Yang is a recently established genus in trib. Tritieeae of Poaceae. In this paper, this genus is taxonomically revised, and a new taxonomic system is presented. The new system includes 3 sections, 26 species and 6 varieties, of which 3 sections and 1 species are described as new, 7 taxa are treated respectively in the ways of new status or combination and reduction. Three new sections, sect. Kengyilia, sect. Stenachyra L. B. Cai and sect. Hyalolepis (Nevski) L. B. Cai, are differentiated by the length and width of spike, the growing position of spikelet, the relative length of glume, the colour of anther and so on. The sect. Kengyilia contains 9 species, distributed in northwestern China, extending westward to Kazakhstan, Kirghizia, Tadzhikistan, Afghanistan and Iran; the sect. Stenachyra L. B. Cai comprises 10 species and 3 verieties, distributed in western China, mainly in the Qinghai-Xizang Plateau; and the sect. Hyalolepis (Nevski) L. B. Cai consists of about 7 species and 3 varieties, distributed in western China, extending westward to Kirghizia and Tadzhikistan. Twenty-six species of Kengyilia are recognized on the basis of the same criterion of specific concept, and the other taxa below the rank of species are also checked in terms of their external morphology. As a result, Kengyilia pen~ dula L. B. Cai is reported as a new species; K. hirsuta var. obviaristata L. B. Cai is raised to a species; K. melanthera (Keng) J. L. Yang, Yen et Baum is reduced to a variety under K. thoroldiana (Oliver) J. L. Yang, Yen et Baum; Roegneria hirsuta var. leiophylla Keng et S. L. Chen is reduced as a synonym of K. hirsuta var. hirsuta; Agropyron thoroldianum var. lasciusculum Melderis is reduced to K. grandiglumis (Keng et S. L. Chen) J. L. Yang, Yen et Baum; Roegneria rigidula var. intermedia Keng et S. L. Chen to K. rigidula (Keng et S. L. Chen) J. L. Yang, Yen et Baum; R. hirsuta var. variabilis Keng et S. L. Chen and R. rnelanthera var. tahopaica Keng et S. L. Chen to K. hirsuta (Keng et S. L. Chen)J. L. Yang, Yen et Baum. In addition, new records onthe geographical distributions of some taxa are also reported in this paper.     

中国禾本科植物的增补

本文发表禾本科中1个新变种,即单穗束尾草;2个新命名,即短芒披碱草与耿耳稃草;17个新组合,即蒙古早熟禾;胎生早熟禾;微药碱茅,疏花以礼草;硬秆草;变种硬秆草;黑药以礼草,大河坝黑药草;梭罗草;窄颖以礼草;孪生以礼草;大颖草;长颖以礼草,糙毛以礼草;无芒以礼草;青海以礼草,阿拉善披碱草。     

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.     

Morphological Identification of Elymus sibiricus,E. nutans,and E. burchan-buddae,and Their Genomic Relationships

In order to investigate morphological variations of Elymus sibiricus L., E. nutans Griseb., and E. burchan-buddae (Nevski) Tzvelev ［=Roegneria nutans (Keng) Keng］, and to explore their systematic relationships, six morphological characters were measured and compared between the Elymus species. Interspecific hybridizations between E. nutans and E. sibiricus, and E. burchan-buddae were carried out with the aid of embryo rescue. Chromosome pairing behaviour was also analysed at metaphase- I of meiosis in pollen mother cells of F₁ hybrids. Morphologic characters, such as length of spikes and length of glumes varied considerably between different individuals of each species, whereas length and width of paleas were quite stable and different between species. The interspecific hybrids were completely sterile and their meioses were irregular. Meiotic configuration of E. nutans × E. burchan-buddae was 7.70I +13.40Ⅱ+0.06Ⅲ+0.08 IV, whereas that of E. sibircus×E. nutans was 11.98I+9.61Ⅱ+0.64Ⅲ+0.39Ⅳ+0.01V. It is concluded from the morphological and cytological study that (1) it is possible to identify the three Elymus species using the palea character, in addition to other traditionally applied characters; (2) Elymus nutans and E. burchan-buddae have a comparatively high genomie relationship, while E. nutans and E. sibircus have a relatively low genomic affinity to one another; and (3) a certain chromosome pairing regulator was presented in the hybrid between E. burchan-buddae andE. nutans.     

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.     

Different maternal genome donor to Kengyilia species inferred from chloroplast trnL-F sequences

To reveal the maternal donor of species in genus Kengyilia, the chloroplast trnL-F sequences of 14 Kengyilia species and several related diploid species were analyzed by using Maximum Parsimony (MP) and Bayesian Inference (BI) methods. The species in Kengyilia were clustered in different clades, which indicated that Agropyron (P) is the likely maternal genome donor to Kengyilia melanthera, K. mutica and K. thoroldiana, while the maternal donor to Kengyilia batalinii, K. nana, K. kokonorica, K. kaschgarica, K. hirsuta, K. alatavica, K. gobicola, K. zhaosuensis, K. rigidula, K. longiglumis and K. grandiglumis was St or Y Roegneria genome.     

Genome constitution of Elymus tangutorum (Poaceae: Triticeae) inferred from meiotic pairing behavior and genomic in situ hybridization

Elymus tangutorum (Nevski) Hand.-Mazz (2n = 6x = 42) is a perennial species in the tribe Triticeae, which distributes in Nepal and north and northwest China. However, the genome constitution of E. tangutorum is controversial and its taxonomic status is not clear. Hybridizations of E. tangutorum were carried out with E. wawawaiensis J. R. Carlson & Barkworth (StH), Roegneria grandis Keng (StY), and E. dahuricus Turcz. ex Griseb. (StYH). Meiotic pairing in the hybrids E. tangutorum × E. wawawaiensis (StH), E. tangutorum × R. grandis (StY), and E. tangutorum × E. dahuricus (StYH) averaged 10.48, 11.12, and 20.92 bivalents per cell, respectively. The results suggested that E. tangutorum is an allohexaploid and contains the StYH genomes. Results of genomic in situ hybridization analysis strongly supported the chromosome pairing data. Therefore, E. tangutorum should be treated as Campeiostachys dahurica var. tangutorum (Nevski) B. R. Baum, J. L. Yang & C. Yen. Intergenomic rearrangements of E. tangutorum may be affected by environmental factors.     

仲彬草属5种植物的核型研究

本文对我国西部高原仲彬草属Ｋｅｎｇｙｉｌｉａ５种植物的核型进行了分析。它们的染色体数目均为２ｎ＝４２,六倍体。核型是：糙毛鹅观草Ｋ．ｈｉｒｓｕｔａ,２ｎ＝６ｘ＝４２＝３６６＋６ｓｍ;青海鹅观草Ｋ．ｋｏｋｏｎｏｒｉｃａ,２ｎ＝６ｘ＝４２＝３６ｍ＋６ｓｍ：黑药鹅观草Ｋ．ｍｅｌａｎｔｈｅｒａ,２ｎ＝６ｘ＝４２＝３８ｍ＋４ｓｍ;硬秆鹅观草Ｋ．ｒｉｇｉｄｕｌａ,２ｎ＝６ｘ＝４２＝３８ｍ＋４ｓｍ;窄颖鹅观草Ｋ．ｓｔｅｎａｃｈｙｒａ,２ｎ＝６ｘ＝４２＝３８ｍ＋４ｓｍ。它们的核型属于１Ｂ或２Ｂ型。染色体中均未发现随体。     

Morphological characteristics of fruits of Fagopyrum (Polygonaceae) from China

Macro- and micro-morphological characteristics of fruits in eight species and one variety of the genus Fagopyrum Mill. (Polygonaceae) from China were observed under stereoscope and scanning electron microscope(SEM). Based on the results, the fruits of the species studied are divided into type Ⅰ , Ⅱ and Ⅲ. The fruits of type Ⅰ are triangular-pyramidal; their surface are rugosely reticulate, neither smooth nor shiny. Two species, F. tataricum and F. dibotrys have this fruit type. Those of type Ⅱ are ovoid-triangular-pyramidal; their surface are smooth and shiny, and striately reticulate. Three species, F. esculentum, F. statice and F. lineare, have this fruit type. In type Ⅲ, the fruits are ovoid-triangular-pyramidal; their surface are smooth and shiny, and covered with many warty grains and sparsely finely striate. F. urophyllum, F. gracilipes, F. leptopodum var. leptopodum, and F. leptopodum var. grossii have this fruit type. Judging from the morphological characteristics of fruits, F. dibotrys might be more closely related to F. tataricumthan to F. esculentum.     

Spore morphology of pteridophytes from China Ⅱ. Sinopteridaceae

Spores of 61 species and 6 varieties in 9 genera of the Sinopteridaceae were examined under scanning electron microscope (SEM). Based on surface ornamentation and other features, the spores of the Sinopteridaceae are divided into three types. In type Ⅰ , the exospore is smooth and the surface ornamentation, which is reticulate, cristate, echinate or rugate, is formed by the perispore. All the other genera of this family, except for Onychium and Cryptogramma, have this pattern of spores. In type Ⅱ, the surface ornamentation is formed by both perispore and exospore. This pattern is found only in Cryptogramma. In type Ⅲ, the perispore is thin and the surface ornamentation is formed by the exospore. Onychium is characterized by this type of spores. Those genera with spores of type Ⅰ of the Sinopteridaceae seem to be closely related to each other and should be natural members of this family. The systematic position of Cryptogramma and Onychium, with spores of type Ⅱ and type Ⅲ respectively, however, should be reconsidered. Aleuritopteris might be the mostprimitive member of the Sinopteridaceae from the evidence of spore morph     

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。     

Phylogenetic relationships in Leymus (Triticeae; Poaceae): Evidence from chloroplast trnH-psbA and mitochondrial coxII intron sequences

Leymus Hochst. is a polyploid genus with a diverse array of morphology, cytology, ecology, and distribution in Triticeae. To investigate the phylogenetic relationships and maternal genome donor of polyploid Leymus, the chloroplast trnH-psbA region and mitochondrial coxII intron sequences of 33 Leymus taxa were analyzed with those of 36 diploid perennial species representing 19 basic genomes in Triticeae. The results showed that reticulate evolution occurred in Leymus species, with the cytoplasmic lineage of Leymus contributed by different progenitors. Interspecific relationships of Leymus were also elucidated on the basis of orthologous comparison. Our data suggested that: (i) due to incomplete lineage sorting and/or difference in the pattern of chloroplast and mitochondrial inheritance, the genealogical conflict between the two genealogical patterns suggest the contribution of Psathyrostachys Nevski, Agropyron J. Gaertn, Eremopyrum (Ledeb.) Jaub. & Spach, Pseudoroegneria (Nevski) Á. Löve, Thinopyrum Á. Löve, and Lophopyrum (Host) Á. Löve to the cytoplasmic lineage of Leymus; (ii) there is a close relationship among Leymus species from the same area or neighboring geographic regions; (iii) L. coreanus (Honda) K. B. Jensen & R. R.-C. Wang, L. duthiei (Stapf) Y. H. Zhou & H. Q. Zhang ex C. Yen, J. L. Yang & B. R. Baum, L. duthiei var. longearistatus (Hack.) Y. H. Zhou & H. Q. Zhang ex C. Yen, J. L. Yang & B. R. Baum, and L. komarovii(Roshev.) C. Yen, J. L. Yang & B. R. Baum are closely related to other Leymus species, and it is reasonable to transfer these species from the genus Hystrix Moench to Leymus; (iv) Leymus species from North America are closely related to L. coreanus from the Russian Far East and L. komarovii from northeast China but are evolutionarily distinct from Leymus species from Central Asia and the Qinghai–Tibet Plateau. The occurrence of multiple origin and introgression could account for the rich diversity and ecological adaptation of Leymus species.     

Comparative anatomy of oil cells and mucilage cells in the leaves of the Lauraceae in China

The distribution density of oil cells, the morphology and structure of both oil and mucilage cells, and their localization in the mesophyll of 112 species, 5 varieties and 2 forms in 21 genera of the Lauraceae are comparatively studied with the methods of tissue clearing and paraffin sectioning. The results show that there exist obvious differences of the distribution density of oil cells among the species in the Lauraceae. The presence of oil cells and mucilage cells is found to be a marked anatomical feature of the leaves in most of the plants in the Lauraceae. Their distribution in the mesophyll can be divided into 4 types: type Ⅰ , in which only oil cells are present; type Ⅱ, in which both the oil and mucilage cells are present; type Ⅲ, in which only mucilage cells are present; type Ⅳ, in which neither oil cells nor mucilage cells are present. The distribution density of oil cells, the distribution types of oil cells and mucilage cells and their localization in the mesophyll are of some taxonomic value at the specific level in the Lauraceae. In the whole Lauraceae or in some large genera, the evolutionary trend of the distribution types of oil cells and mucilage cells might be as follows: type Ⅰ → type Ⅱ →type Ⅲ →type Ⅳ. The characteristics of the 4 distribution types of oilcells and mucilage cells support the division of two subfamilies in the Lauraceae.     

内蒙古冰草属（禾本科）一新变种——毛稃沙芦草

报道了内蒙古冰草属一新变种——毛稃沙芦草(Agropyron mongolicum Keng var. helinicum L.Q.Zhao et J. Yang)。该变种外稃密被长柔毛,颖光滑无毛不同于沙芦草（A. mongolicum Keng）和毛沙芦草（A. mongolicum Keng var. villosum H.L.Ying）。     

Intergenomic rearrangements after polyploidization of Kengyilia thoroldiana (Poaceae: Triticeae) affected by environmental factors

Polyploidization is a major evolutionary process. Approximately 70–75% species of Triticeae (Poaceae) are polyploids, involving 23 genomes. To investigate intergenomic rearrangements after polyploidization of Triticeae species and to determine the effects of environmental factors on them, nine populations of a typical polyploid Triticeae species, Kengyilia thoroldiana (Keng) J.L.Yang et al. (2n = 6x = 42, StStPPYY), collected from different environments, were studied using genome in situ hybridization (GISH). We found that intergenomic rearrangements occurred between the relatively large P genome and the small genomes, St (8.15%) and Y (22.22%), in polyploid species via various types of translocations compared to their diploid progenitors. However, no translocation was found between the relatively small St and Y chromosomes. Environmental factors may affect rearrangements among the three genomes. Chromosome translocations were significantly more frequent in populations from cold alpine and grassland environments than in populations from valley and lake-basin habitats (P<0.05). The relationship between types of chromosome translocations and altitude was significant (r = 0.809, P<0.01). Intergenomic rearrangements associated with environmental factors and genetic differentiation of a single basic genome should be considered as equally important genetic processes during species'' ecotype evolution.     

鹅观草属4个种的核型与进化

报道了4个种鹅观草属（Roegneria C. Koch）植物的核型,其核型公式如下：芒颖鹅观草(R. aristiglumis Keng et S. L. Chen), 2n=4x=28=22m+6sm（2SAT）;短颖鹅观草(R. breviglumis Keng), 2n=4x=28=20m（2SAT）+8sm;红原鹅观草(R. hongyuanensis L. B. Cai), 2n=4x=28=22m+6sm（2SAT）;山东鹅观草(R. shandongensis（B. Salomon）J. L. Yang, Y. H. Zhou et Yen), 2n=4x=28=24m（2SAT）+4sm。同时对染色体主要特征的演变规律进行了分析,揭示了鹅观草属4个种的相对进化程度以及宏观分类中2个组的系统发育关系,表明鹅观草属的半颖组在系统发育中派生了颖体短小的小颖组。     

Cytogenetic analyses inKengyilia laxiflora (Poaceae, Triticeae)

Kengyilia laxiflora (2n = 42) was cytogenetically studied with testersK. hirsuta (2n = 42, PPStStYY) andRoegneria kamoji (2n = 42, HHStStYY). Our data suggested thatK. laxiflora may possess a modified P genome, the P¹. Its St and Y genomes may also contain some structural changes and are more closely related to the St and Y genomes ofK. hirsuta than to those ofR. kamoji. Kengyilia laxiflora is reproductively separated from bothK. hirsuta andR. kamoji. These results indicated thatK. laxiflora is a good species in the genusKengyilia.     

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。     

用RAPD分子标记探讨鹅观草属的种间关系

通过34个10碱基随机引物对鹅观草属(RoegneriaC.Koch)26个物种进行PCR扩增,28个引物能产生多态带。在NTSYS程序中,对16个引物扩增的186条扩增产物,计算Jaccard遗传相似系数,建立UPGMA聚类图。结果表明:(1)物种间遗传差异明显,具有丰富的遗传多样性;(2)StY和StYH基因组的物种存在着一定的遗传差异,并各有一定程度的分化,这种分化是与地理位置相联系的。相距越远,物种相似程度越低;(3)形态差异较小,基因组同源,地理分布一致的物种聚类在一起,表现出较密切的亲缘关系;(4)无芒类群的R.alashanicaKeng和R.magnicaespes(D.F.Cui)L.B.Cai与其余分析的24个Roegneria物种存在着极大的遗传差异;(5)分布于西亚的R.caucasicaC.Koch.与分布于东亚和中亚的物种存在着较大的RAPD变异,亲缘关系较远;(6)在对R.ciliaris(Trin)Nevski和R.japonensis(Honda)Keng、R.tenuispicaJ.L.YangetY.H.Zhou和R.pendulinaNevski、R.tsukushiensis(Honda)Ohwi和R.kamojiOhwi等的分类处理上,基本上与形态学和细胞学的研究结果一致。对RAPD标记在Roegneria系统分类研究中的应用进行了讨论。