杉科的细胞分类学和系统演化研究

根据杉科的核型资料,本文(1)提出“1B”可能是一个新的高等植物核型类型;(2)讨论了各属的有关分类学问题及相互亲缘关系,它们的进化顺序可能是柳杉属、水松属、落羽杉属、水杉属、巨杉属、红杉属、杉木属(密叶杉属与之近缘)、台湾杉属;(3)支持金松属分立成金松科,它可能比杉科各属原始;(4)红杉(AAAABB)的亲本可能是二个古代种水杉”和“巨杉”,它们的直接后裔或留下的近缘是水杉和巨杉;(5)杉科存在A和L两条进化路线,前者包括柳杉属、水松属、落羽杉属、台湾杉属,后者包括水杉属、巨杉属、红杉属、杉木属(密叶杉属);(6)提出一个杉科新系统(包括一个新亚科)：I．柳杉亚科(柳杉属),II．落羽杉亚科(水松属、落羽杉属),III．红杉亚科(冰杉属、巨杉属、红杉属),Ⅳ.杉木亚科(杉木属、密叶杉属),V．台湾杉亚科,新亚科(台湾杉属)。本文还对前人的杉科系统作了讨论。     

杉科的细胞分类学和系统演化研究

根据杉科的核型资料,本文(1)提出“1B”可能是一个新的高等植物核型类型;(2)讨论了各属的有关分类学问题及相互亲缘关系,它们的进化顺序可能是柳杉属、水松属、落羽杉属、水杉属、巨杉属、红杉属、杉木属(密叶杉属与之近缘)、台湾杉属;(3)支持金松属分立成金松科,它可能比杉科各属原始; (4)红杉(AAAABB)的亲本可能是二个古代种“水杉”和“巨杉”,它们的直接后裔或留下的近缘是水杉和巨杉;(5)杉科存在A和L两条进化路线,前者包括柳杉属、水松属、落羽杉属、台湾杉属;后者包括水杉属、巨杉属、红杉属、杉木属(密叶杉属);(6)提出一个杉科新系统(包括一个新亚科):Ⅰ.柳杉亚科(柳杉属),Ⅱ.落羽杉亚科(水松属、落羽杉属),Ⅲ.红杉亚科(水杉属、巨杉属、红杉属),Ⅳ.杉木亚科(杉木属、密叶杉属),Ⅴ.台湾杉亚科,新亚科(台湾杉属)。本文还对前人的杉科系统作了讨论。     

生境的破坏或改变对杉科种属分布的影响

杉科共有10个属16种,其中5个是单种属(水杉属、水松属、巨杉属、北美红杉属、金松属),5个是寡种属(杉木属、台湾杉属、柳杉属、落羽杉属、密叶杉属),两者各占50%。分布于北温带地区,多在东亚和北美洲,仅密叶杉属分布于南半球澳大利亚的塔斯马尼亚。植物与生境是一个互相依存的统一体,而生境的破坏或改变对杉科植物种属分布有严重的影响。在美国加利福尼亚洲现今仅有巨杉和北美红杉两个单种属,这是由于火山爆发留下来,现在仅分布于美国加州部分地区。     

杉科的两条演化路线

本文根据核型资料提出杉科可能存在A、L两条演化路线,前者包括由原始到进化的柳杉属、水 松属、落羽杉属、台湾杉属,以平均臂比快速增加、染色体长度比缓慢增加为特征;后者包括依序进化的水杉属、巨杉属、红杉属、杉木属(可能还有密叶杉属),以平均臂比缓慢增加、染色体长度比迅速增加为特点。该结论也得到形态学、解剖学、胚胎学等资料的支持。     

密叶杉属的核型分析及其系统位置的探讨

本文首次对特产澳大利亚塔斯马尼亚岛的Athrotaxis cupressoides和Athrotaxis selaginoides进行了核型分析,核型公式分别为2n=22=22m(2SAT)和2n=22=20(2SAT)+2sm,均属Stebbins的1B类型,它们的染色体相对长度组成为22=2L+10M₂+8M_l+2S和22=2L十10M₂+6M_l+4S,后者比前者较为进化。根据密叶杉属和杉科其他各属核型资料的比较分析,它们由原始到进步的顺序可能为：柳杉属、水松属、落羽杉属、水杉属、巨杉属、红杉属,密叶杉属、杉木属和台湾杉属。 密叶属与红杉属、巨杉属和杉木属较为近缘。 这些在以染色体长度比和平均臂比为纵、横坐标的图上得到清楚反映。 根据核型资料,密叶杉属以隶于单型的亚科Arthrotaxoideae较为合适,这也得到形态学、胚胎学、孢粉学和地理学资料的支持。 本文还对前人系统中的密叶杉属位置进行了讨论。     

中国和邻近地区杉科特有植物的核型及其进化水平的初步研究

中国和邻近地区杉科特有植物的核型类型,柳杉属、水松属、水杉属均为“1A”,杉木属、台湾杉属分别为“1B”和“2B”,它们的进化水平似依序递增并组成A-柳杉属——水松属——台湾杉属和L-水杉属——杉木属二条演化路线。该结果得到某些形态学性状和古植物学的支持,基本上适用于整个杉科。     

杉科植物的起源、演化及其分布

本文根据对杉科的系统发育、现代分布和化石分布的研究,结合古地理和古气候资料,讨论了杉科的起源、演化和现代分布格局的成因。杉科基本上是一个亚热带科,我国长江、秦岭以南至华南一带是其现代分布中心。东亚中高纬度的东北地区可能是其起源中心和早期分化中心。起源时间为早侏罗纪或晚三叠纪。杉科植物的各种类型很可能在早白垩纪甚至晚侏罗纪就已分化出来。杉科植物于东亚起源后,在当时劳亚古陆尚未完全解体、气候分带现象尚不甚明显的情况下跨越欧亚大陆散布到北美,并扩散到南半球。自晚白垩纪,白令陆桥和北大西洋陆桥对其在北半球的散布发挥了重要作用。杉科植物目前虽处于衰退状态,但在地质史上却曾经经历过极其繁盛的时代。在中生代中晚期和早第三纪,杉科植物种类繁多,广布于北半球,向北扩散到北极圈内的高纬度地区,是当时的大科。大多数现存属曾分别有过3个或2个分布中心：水松属、落羽杉属和北美红杉属在东亚、北美西部和欧洲;水杉属在东亚和北美西部;柳杉属、杉木属,很可能也包括台湾杉属在东亚和欧洲;巨杉属在欧洲、北美和东亚。在晚白垩纪和第三纪,现存属特别是水松属、落羽杉属、水杉属、北美红杉属和巨杉属,曾是北半球森林植被的重要组成成分。南半球也曾有少量种类,分布亦远较现代普遍。杉科在白垩纪的多样性达到鼎盛,具所有的现代属和大量的化石器官属,但在以后漫长的历史发展过程中,由于地质变迁、气候变化,大量类群绝灭。晚第三纪全球性的气温下降迫使杉科逐渐从高纬度地区撤出。第四纪冰期气候的剧烈恶化使杉科分布区进一步显著退缩至中、低纬度地区,最后在欧洲全部消失,仅在东亚、北美及澳大利亚的少数几个植物 “避难所”中残存下来。现今各属多分布于环太平洋地区极为狭窄的局部范围,在分布区内呈现出孤立或星散的残遗分布式样。杉科现存各属均为古老的孑遗或残遗类群。     

秃杉的胚胎发育及其与杉科各属的比较

本文报道秃杉(Taiwania flousiana Gaussen)的雌雄配子体和胚胎发育的研究结果及其与 杉科各属的比较。秃杉的大孢子经10次游离核同时分裂,产生1019—1021个游离核。 6月 初游离核开始向心地形成细胞壁。雄配子体具两个大小、形状相同的精子。受精作用在湖北 发生于6月底至7月初(1960)。受精卵经3次游离核分裂,形成8核原胚,排列成两群,紧接 着形成壁。上下2群细胞的数目分配为5：3或4:4,少数为6:2。 从胚胎学的资料来看,秃杉与台湾杉一样,它们最接近杉木属和水松属,与柳杉属、落羽松 属和水杉属也有类似之处。它们的共同特征是：精子形状、大小相同,复合颈卵器顶生,原胚 具8个游离核,兼具裂生多胚。而红杉属、巨杉属、密叶杉属和金松属的胚胎发育各有特点,应分别属于杉科不同的类群。     

杉科植物的分类学研究

本文对杉科植物的研究历史作了回顾,根据分支分类结果和表征分类结果,提出了一个新的分类系统.以形态学为依据,结合其他学科的研究成果,对杉科的分类作了订正。作者承认杉科植物共9属、12种及3变种,将杉木、厚皮杉木、德昌杉木和米德杉木归并,支持柳杉作为日本柳杉的变种、台湾杉木作为杉木的变种、秃杉和台湾杉归并的观点。     

杉科植物分类学研究

本文对杉科植物的研究历史作了回顾,根据分支分类结果和表征分类结果,提出了一个新的分类系统。以形态学为依据,结合其他学科的研究成果,对杉科的分类作了订正。作者承认杉科植物共９属、１２种及３变种,将杉木、厚皮杉木、德昌杉木和米德杉木归并,支持柳杉作为日本柳杉的变种、台湾杉木作为杉木的变种、秃杉和台湾杉归并的观点。     

Phylogenetic Analysis of the Family Taxodiaceae

In the present paper,both cladistic analysis and phenetic analysis were conducted to evaluate the phylogenetic relationships of the Taxodiaceae based on an extensive literature review and study of herbarium． In the cladistic analysis,the Sciadopityaceae was chosen as outgroup．The polarity of characters was determined mainly according to outgroup comparison,fossil evidence and generally accepted viewpoints of morphological evolution．By the result of compatibility analysis,character 2(leaf type),which possessed a much higher coefficient than others whether or not its polarity was altered,was deleted． Finally,a data matrix consisting of all the extant nine genera and 24 characters was analyzed using Maximal Same Step Method,Synthetic Method,Evolutionary Extremal Aggregation Method and Minimal Parallel Evolutionary Method,and four cladograms were generated,of which only the most parsimonious one (Fig．1)was presented for discussion. The cladogram shows that the Taxodiaceae are assorted along five lines of evolution： 1)Metasequoia;2)Sequoiadendron,Sequoia;3)Cryptomeria;4)Glyptostrobus and Taxodium;5)Cunninghamia,Athrotaxis and Taiwania． Ten genera(including Sciadopitys)and 59 characters were used in the phenetic analysis．The phenogram(Fig．2)indicates that Sciadopitys is a very distinct group with remote affinity to the other genera,and the Taxodiaceae are divided into four groups：1)Sequoia,Sequoiadendron;2)Athrotaxis,Cunninghamia and Taiwania;3)Cryptomeria,Glyptostrobus and Taxodium;4)Metasequoia． Based primarily on the result of cladistics,with reference to that of phenetics,the main conclusions were drawn as follows：(1)Generic relationships：Cryptomeria should be considered the most primitive genus in the extant groups of the Taxodiaceae. Glyptostrobus and Taxodium, close to Cryptomeria, are sister taxa and relatively primitive groups. Sequoiadendron and Sequoia are closely related and intermediate advanced. Metasequoia is a more or less isolated taxon, relatively close to Sequoiadendron and Sequoia. Cunninghamia. Athrotaxis and Taiwania might represent a single lineage and form a very advanced group, of which Taiwania may be the most specialized. (2) Systematic treatments: The authors support the viewpoint that Sciadopitys should be treated as an independent family, and suggest that the Taxodiaeae should be divided into five tribes. Systematic arrangements are as follows: Taxodiaceae Warming Trib. 1. Cryptomerieae Vierhapper Gen. 5. Sequoia Endl. Gen. 1. Cryptomeria D. Don Trib. 4. Metasequoieae Pilger et Melchior Trib. 2. Taxodieae Benth. et Hook. Gen. 6. Metasequoia Miki ex Hu et Cheng Gen. 2. Glyptostrobus Endl. Trib. 5. Cunninghamieae Zucc. Gen. 3. Taxodium Rich. Gen. 7. Cunninghamia R. Br. Trib. 3. Sequoieae Wettstein Gen. 8. Athrotaxis D. Don Gen. 4. Sequoiadendron Buchholz Gen. 9. Taiwania Hayata     

The Karyotype Analysis of Arthrotaxis (Taxodiaceae) and Its Systematic Position

The present paper deals for the first time with an analysis of the karyotypes of Athrotaxis cupressoides Don and A. selaginoides Don endemic to Tasmania (Australia). Their morphology of somatic chromosomes in seed root-tip cells, chromosome measurements, and diagrams are shown in Plate 1, Table 1 and Fig. 1 respectively, The karyotypic formulas of the two species are 2n = 22 = 22m (2SAT) and 2n = 22 = 20m(2SAT ) + 2sm according to of terminology Lexvan et al (1964). They all belong to IB type of Stebbins’(1971)karyotypic asymmetry which was reported for the first time in the higher plants by Li(1987b). Their chromosome complements are 22 = 2L + 10M₂+ 8M₁+ 2S and 22 =2L+ 10M₂+ 6M₁+ 4S respectively according to the standard defined by Kuo et al. (1972) based on relative length. The karyotype of A. selaginoides is more advanced than that of A. cupressoides. In the light of karyotypic data, the sequence of the taxo-diaceous genera (excl. Sciadopitys) from primitive to advanced may be in the following order: Cryptomeria, Glyptostrobus, Taxodium, Metasequoia, Sequoiadendron, Sequoia, Athrotaxis, Cunninghamia and Taiwania. The genus Athrotaxis is closely related to Sequoia (Sequoiadendron) and Cunninghamia The peculiarity of the karyotype of Athrotaxis deserves the establishment of a new status Arthrotaxoideae (Wettstein) L. C. Li This suggestion is also supported by the data from morphology, embryology, palynology and geography. The family Taxodiaceae is divided into six subfamilies and nine genera, as shown in the following table:————————————————————————————————————————————————— 1. Cryptomerioideae Hida Cryptomeria D. Don 4. Arthrotaxoideae (Wettstein) L. C. Li 2. Taxodioideae Pilger Glyptostrobus Endl. Taxodim Richard Athrotaxis D. Don 3. Sequoideae Saxton metasequoia Miki ex Hu et 5. Cunninghamioideae Hida Cunninghamia Cheng Sequoiadebron Buch. Sequoia Endl. R.Brown 6. Taiwanioideae (Hayata)L. C. Li Taiwanta Hayata————————————————————————————————————————————————— The systematic positions of Athrotaxis in the systems of other authors are dis-cussed too.     

杉科、柏科的系统发生分析——来自28S rDNA序列分析的证据

对杉科（Taxodiaceae）与柏科（Cupressaceae s.s.）的28S rRNA基因的部分序列（约630 bp）进行PCR扩增、序列测定和系统发生关系分析,用简约法和邻接法构建的系统发生树基本一致。结果表明,杉科与柏科构成一个单系群,支持将杉科、柏科（Sciadopitys除外）合并为一个科——广义柏科（Cupressaceae sensu lato）的观点。在广义柏科中,Taiwania、Athrotaxis分别形成一支系;Metasequoia、Sequoia、Sequoiadendron关系较近,聚成一支系; Taxodium、Glyptostrobus、Cryptomeria聚成一支系;柏科聚成一支系。这一分析结果与叶绿体基因序列的分析结果相吻合,但是由于28S rRNA基因的进化速率较慢,尚不能分辨上述各个支系之间的系统演化关系。 Abstract:DNA sequences from 28S rDNA were used to assess relationships between and within traditional Taxodiaceae and Cupressaceae s.s.The MP tree and NJ tree generally are similar to one another.The results show that Taxodiaceae and Cupressaceae s.s.form a monophyletic conifer lineage excluding Sciadopitys.In the Taxodiaceae-Cupressaceae s.s.monophyletic group,the Taxodiaceae is paraphyletic.Taxodium,Glyptostrobus and Cryptomeria forming a clade(Taxodioideae),in which Glyptostrobus and Taxodium are closely related and sister to Cryptomeria;Sequoia,Sequoiadendron and Metasequoia are closely related to each other,forming another clade (Sequoioideae),in which Sequoia and Sequoiadendron are closely related and sister to Metasequoia;the seven genera of Cupressaceae s.s.are found to be closely related to form a monophyletic lineage (Cupressoideae).These results are basically similar to analyses from chloroplast gene data.But the relationships among Taiwania,Sequoioideae,Taxodioideae,and Cupressoideae remain unclear because of the slow evolution rate of 28S rDNA,which might best be answered by sequencing more rapidly evolving nuclear genes.     

Two Evolutionary Lines of Taxodiaceae

According to the karyotypic data the present author proposes two evolutionary lines, A and L (Fig. 1), in Taxodiaceae (Exclud. Sciadopitys). The former is characterized by a relatively rapid increase of the mean arm ratio but a relatively slow rise of the ratio of the longest chromosome to the shortest one and it is composed of Cryptomeria, Glyptostrobus, Taxodium and Taiwania, which advance from primitive to progressive in the order. The latter is characterized by, on the contrary, a relatively slow increase of mean arm ratio and a relatively rapid rise of the ratio of chromosome size and it comprises Metasequoia, Sequoiadendron, Sequoia and Cunninghamia (probably Athrotaxis also), which advance in the order. The inference is supported by the data from morphology, anatomy, embryology and so on.Key words Taxodiaceae; Karyotype; Evolutionary line     

Phylogenetic relationships in Taxodiaceae and Cupressaceae sensu stricto based on matK gene, chlL gene, trnL-trnF IGS region, and trnL intron sequences

Nucleotide sequences from four chloroplast genes, the matK, chlL, intergenic spacer (IGS) region between trnL and trnF, and an intron of trnL, were determined from all species of Taxodiaceae and five species of Cupressaceae sensu stricto (s.s.). Phylogenetic trees were constructed using the maximum parsimony and the neighbor-joining methods with Cunninghamia as an outgroup. These analyses provided greater resolution of relationships among genera and higher bootstrap supports for clades compared to previous analyses. Results indicate that Taiwania diverged first, and then Athrotaxis diverged from the remaining genera. Metasequoia, Sequoia, and Sequoiadendron form a clade. Taxodium and Glyptostrobus form a clade, which is the sister to Cryptomeria. Cupressaceae s.s. are derived from within Taxodiaceae, being the most closely related to the Cryptomeria/Taxodium/Glyptostrobus clade. These relationships are consistent with previous morphological groupings and the analyses of molecular data. In addition, we found acceleration of evolutionary rates in Cupressaceae s.s. Possible causes for the acceleration are discussed.     

杉科的染色体资料及在系统演化研究中的作用

杉科共由10属(包括金松属)、20种(变种)组成。本文整理了19种(占95％,隶10属)植物的染色体数目和16种(占80％,隶9属)的核型资料,核型的模式图如图1所示。通过对这些细胞学资料的分析,笔者支持2n=20、x=10的金松属从杉科(2n=22、x=11)分立成金松科。根据其他各属间的亲缘关系,本作者认为可以把它们分隶于5个亚科:Ⅰ.柳杉亚科:Cryptomerioideae(Cryptomeria);Ⅱ.落羽杉亚科Taxodioideae(Glyptostroous,Taxodium);Ⅲ.红杉亚科Sequoideae(Metasequoia,Sequoiadendron,Sequoia);Ⅳ.杉木亚科Cunninghamioideae(Cunninghamia,还可能有:Athrotaxis);Ⅴ.台湾杉亚科Taiwainoideae(Taiwania)。这些亚科和属的进化水平依序渐增,它们分别位于进化路线A(亚科Ⅰ、Ⅱ、Ⅴ)和进化路线L(亚科Ⅲ、Ⅳ)上。这些结果是前人的演化系统所没有涉及的,表明了染色体资料在杉科的系统演化研究中起着重要的作用。     

PROANTHOCYANIDINS IN NEEDLES FROM SIX GENERA OF THE TAXODIACEAE

Proanthocyanidin contents of needles ranged from a mean of 150 to 300 μg per mg dry wt in five species from five genera of the Taxodiaceae, Sequoiadendron giganteum (Lindl.) Bucch., Metasequoiaglyptostroboides H. Hu and Cheng, Sequoia sempervirens (D. Don) Endl., Taxodium distichum L. Rich., and Sciadopitys verticillata Siebold and Zucc. However, significantly lower amounts (70 μg per mg dry wt) were found in Cryptomeria japonica (L.f.) D. Don. This latter species as well as Sciadopitys verticillata, contained little or no prodelphinidin, while the other four species contained a ratio of procyanidin to prodelphinidin up to about 1:5. These data were based on analyses from three trees from each species. In addition, one tree from each species was examined in more detail. The major flavan-3-ol in all cases was (+)-catechin, with only non-detectable or trace amounts of (–)-epicatechin. The triphenolic flavan-3-ol, (+)-gallocatechin, was a minor constituent in all species, except Sciadopitys and Cryptomeria. The (–)-epigallocatechin was detected in Metasequoia, Sequoiadendron and Sequoia. All contained either (–)-epicatechin-(+)-catechin or (+)-catechin-(+)-catechin as the major procyanidin dimer. Prodelphinidin dimers were only tentatively identified.     

从木材解剖特征初拟针叶树分类系统

<正> 针叶树(Conifers)包括银杏纲和松杉纲的树种。根据《中国植物志》第七卷记载,现代裸子植物的种类分属于4纲9目12科71属约800种,我国产4纲8目11科41属236种,栽培1科7属51种。从国产针叶树材的解剖特征观察,银杏纲木材构造简单,仅具管胞、含晶细胞和射线细胞。管胞排列不整齐,射线薄壁细胞垂直壁无节状加厚,四隅无凹痕,轴向薄壁细胞缺,射线仅单例,所以是针叶村最原始的一个纲,应排列在松杉纲之前。     

On the Ultrastructure of Pollen Exine in Metasequoia Hu and Cheng

Metasequoia is endemic to China. Present study deals with ultrastructure of pollen exine of M. glyptostroboides Hu et Cheng, and in comparision with other genera of the family. Pollen grains of Metasequoia are spheroidal or subsphoroidal and 27.8(24.3–32.3) μm in diameter. There is a papilla in the distal face. The papilla is wide at the base, 3.5–5.2 μm high, with pointed and circular end and the base crooked toward one side. Exine is about L5 μm thick, layers distinct, Nexine is as thick as sexine. Surface weakly granulate. According to observation by SEM, exine is covered with fine granules and rather coarse tuberculae. The former can be easily separated from the latter. The loose and uneven tuberculae are provided with minute spinules on the surface and generally fall off after acetolysis. The fine and dense granulae, however, remain intact after acetolysis. The study by TEM shows that ektexine is made of granules densely arranged and connected with each other. In addition, sparse Ubisch bodies are unevenly distributed on granular layer with geminate surface. The thick endexine, is composed of 10–15 lamellae. It is worthy to note that all lamellae possess tripartite structure. But lamellae of endexine in other genera of Taxodiaceae have no tripartite structure except the lamella near ektexine. Number of lamella and thickness of endexine in Metasequoia differ from those of other genera in Taxodiaceae; for example endexine with 8–10 lamellae in Taxodium, 8–9 lamellae in Sequoia, 6–7 lamellae in Glyptostrobus, 6–8 lamellae in Cunninghamia, about 16 lamellae in Cryptomeria etc.