侧柏雌球果及其胚珠的发育

观察了侧柏（Ｐｌａｔｙｃｌａｄｕｓ ｏｒｉｅｎｔａｌｉｓ（Ｌ．）Ｆｒａｎｃｅ）胚珠的发育过程及后期球果苞片的结构变化。在北京,雌球果原基７月分化。通常一个球果有４对苞片,中部两对可育,靠球果顶端一对各产生一枚胚珠,其下一对各两枚。胚珠的发育顺序是向顶的,下部可育苞片腑部的两枚胚珠源于同一原基。胚珠原基分化成珠心和珠被,在发育过程中,珠被逐渐包围珠心,最后形成２烧瓶状的胚珠。１１月至次年１月,球果处     

侧柏雌球果及其胚珠的发育

Seed cone in Platycladus orientalis (L.) France consists of four or five pairs of decussate bracts. Usually, two pairs of the fertile bracts in the middle of the cone subtend six ovules, which initiate in an acropetal manner. Only one ovule presents on each of the upper fertile bract, while two ovules initiate from a common primordium in the axil of lower bracts. In Beijing, most female cones initiated in July. All parts of the cone formed before dormancy, which occurred during November to the next January. After pollination in March, bract morphology changed dramatically; intercalary growth of the bract base formed a conspicuous protuberance, in which inverted vascular system developed. Furthermore, ovules on different pairs of bracts initiated in an acropetal manner and two ovules in each lower fertile bract initiated from a common primordium, which was different from the basipetal initiation of ovules and independently formed single ovule as reported by Takaso in Calltris.     

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     

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.     

CONE AND OVULE ONTOGENY IN TAXODIUM AND GLYPTOSTROBUS (TAXODIACEAE – CONIFERALES)

Seed cones in Taxodium distichum and Glyptostrobus pensilis occupy the position of permanent shoots and are initiated in the summer preceding spring pollination. Morphological features are similar in the two genera, reflecting their close taxonomic relationship. Ovule complexes originate as two (rarely more) ovule primordia in the axil of each fertile bract but without any indication of a preceding discrete ovuliferous scale. When the nucellus, integument, and micropyle are well developed, a series of up to ll abaxial lobes forms at the base of each ovule pair. They become fused by basal growth. After pollination the common basal meristem of lobes and bract extends by intercalary growth to form the conspicuous “ovuliferous scale” of the mature cone; the lobes enlarge and exceed the ovules. Despite the topographic similarity in the cones of both genera, there are differences in vasculature such that the vascular traces to the axillary complex originate directly from the axial cylinder in Glyptostrobus but from the bract trace in Taxodium. The complex vasculature of the mature cone develops late and primarily as an expression of intercalary growth.     

杉科植物的系统发育分析

本文以形态学为依据,参考其他学科的研究成果,用分支分类方法并结合表征分类方法探讨了杉科植物的系统演化关系,提出了新的分类系统。在分支分类中,金松科被选作外类群。主要根据外类群比较原则、化石原则和一般的演化规律,确定了性状的祖征和衍征,采用最大同步法、综合分析法、演化极端结合法及最小平行进化法共四种方法进行分支分析,选择最简约的分支图作为本文讨论基础。在表征分类中,选取59个性状,利用距离系数和类平均法,对金松属和杉科各属进行了聚类运算,得出表征图。综合两种分析结果,主要结论如下:(1)属间关系:柳杉属是现存杉科植物中最原始的类群。水松属和落羽杉属关系密切,二者与柳杉属近缘。巨杉属和北美红杉属关系密切,是中级进化水平的类群。水杉属与巨杉属和北美红杉属的亲缘关系相对较近。杉木属、密叶杉属和台湾杉属关系密切,是杉科植物中的高级进化类群,其中又以台湾杉属演化水平最高。(2)系统排列:支持金松科的成立,将杉科分成5族,即柳杉族(仅含柳杉属)、落羽杉族(含水松属、落羽杉属)、北美红杉族(含巨杉属、北美红杉属)、水杉族(仅含水杉属)和杉木族(含杉木属、密叶杉属及台湾杉属)。     

Cone and ovule development in Cunninghamia and Taiwania (Cupressaceae sensu lato) and its significance for conifer evolution

We examined the early developmental stages of the seed cones and seeds of two conifer genera, Cunninghamia and Taiwania, using scanning electron microscope (SEM) images of freshly collected material. In recent similar studies, these two taxa were not described. The present paper aims to fill that gap. Both genera appear to have features crucial to the understanding of the evolution of the cupressaceous cone, characteristic of the families Cupressaceae and Taxodiaceae, and provide further evidence for the need to merge these families. These features are: the ovuliferous scale in Cunninghamia develops as a small lobe with each of three ovules; in Taiwania these lobes are absent, but a small ridge could be a vestige of them. In neither of these two genera does an ovuliferous scale develop to maturity and only limited intercalary growth transforms the bracts, of which only their width and final shape distinguishes them from sterile leaves. Thus, the bracts, not the ovuliferous scales, form the mature cone in these two genera. This trend is continued in more derived genera of Cupressaceae. Another key extant taxon that has helped to elucidate the evolution of this type of conifer cone is Sciadopitys; similar studies have already been done on this genus, and we compared our findings to them. We also considered certain fossil Mesozoic conifer cones, which shed further light on the evolution of the cupressaceous cone. The evidence from these various genera strongly indicates that recently reconstructed phylogenies of gymnosperms based on molecular evidence from extant taxa do not reflect the evolution that actually happened. Such studies need to take into account nonmolecular evidence, as detailed here.     

从形态学看北美红杉的亲本

本文通过形态性状的比较研究认为AAAABB的同源异源多倍体红杉可能由亲本水杉(属)和巨杉(属)杂交起源,它们分别为红杉提供了染色体组AA和染色体组B。这与根据核型等资料的分析结果相吻合。但跟Stebbins认为红杉除水杉(属)以外的另一个亲本已经灭绝并没有留下近缘的推论存在分歧。     

水杉属和红杉属化石叶表皮鉴定参照系的特殊性

杉科植物的许多属种在小枝的形态和叶片排列上相似,而杉科植物的化石标本多保存为枝叶形式。表皮的特征作为压型化石枝叶标本细胞信息的重要来源,甚至是惟一来源。一直作为杉科植物化石分类鉴定的主要依据。水杉和北美红杉分别是水杉属和红杉属植物化石的惟一现存最近亲缘种,以往关于北美红杉的气孔分布和排列等方面的报道存在分歧,根据作者的研究,北美红杉的表皮特征变异幅度非常广。水杉的气孔分布也与以往报道有差异。利用表皮的特征鉴定杉科植物化石时;不同的处理方法和处理时间,角质层的完整程度和观察数量等均可以影响植物表皮特征的正确获取。     

从核型看北美红杉的起源

本文分析了同源异源自然六倍体北美红杉的AA和B染色体组的组型,它们属Stebbins的“1A”和“1B”,经与杉科各属植物的核型比较后发现分别与相同类型的水杉和巨杉最为接近,因而认为水杉属和巨杉属的某个古代种可能是北美红杉的二个杂交亲本,水杉和巨杉则或许是它们的后裔或近缘。本研究支持Stebbins认为水杉属的某个古代种是红杉的一个祖先而水杉是它的直接后裔的假设,但与他认为另一个祖先已经灭绝并没有留下近缘的推测存在分歧。     

Morphology, Anatomy and Ontogeny of Female Cones in Acmopyle pancheri(Brongn. & Gris) Pilg. (Podocarpaceae)

The developmental morphology and anatomy of the female conesof Acmopyle pancheri(Brongn. & Gris) Pilg. (Podocarpaceae)are described and illustrated, based on observations, histology,scanning electron microscopy (SEM) and nuclear magnetic resonance(NMR) imaging. Ovulate development is typically podocarpaceous.Ovules are unitegmic, and horizontal or inclined upwards throughoutontogeny; the pollination drop is inverted because of the declinatemicropyle. Ontogeny of the epimatium-ovule complex is acropetal,the epimatium developing first. A terminal, distal sterile bractcreates a pollen-scavenging area. During development, the wholecone re-orientates through some 270°, and the seed realignsapprox. 60° with respect to the receptacle axis. The ‘receptacle’or podocarpium supporting the seed is formed by gradual fusionof initially free bracts. The structures adnate to these bractsrepresent homologues of ovuliferous scales; they bear vestigialepimatia which may develop into supernumerary ovules or non-functionalepimatia. Thus, female cones ofA. pancheri are vestigially multi-ovulate.NMR imaging effectively and non-invasively revealed the three-dimensionalarrangement of vascular bundles and resin canals in the cones.Copyright 2001 Annals of Botany Company Acmopyle pancheri(Brongn. & Gris) Pilg., anatomy, developmental morphology, gymnosperms, histology, nuclear magnetic resonance (NMR) imaging, ontogeny, ovules, Podocarpaceae, scanning electron microscopy (SEM), seed cones     

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.     

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.     

CONE DEVELOPMENT IN LIBOCEDRUS (CUPRESSACEAE)—PHENOLOGICAL AND MORPHOLOGICAL ASPECTS

Cones in Libocedrus plumosa are initiated in New Zealand in February (late summer) and continue to develop through winter. The ultimate pair of bracts protrude and continue extension growth while the apex remains flat. Ovules are initiated in two pairs in July on the cone apex (i.e., are axial in origin), alternate with the upper two pairs of bracts. Ovule differentiation proceeds rapidly to the stage of pollination; the cone apex may develop further as a short columella. Soon after ovule initiation an adaxial ligulelike outgrowth of each member of the upper two pairs of bracts is initiated, that of the ultimate pair being broader than that of the lower pair. By intercalary growth of each ligule base an enlarged structure is developed, displacing the bract into a lateral position and forming the four scales that enclose the developing seeds in a valvate manner. Cone and seed maturation follows with seed dispersal in March, approximately 1 year after cone initiation. The results show that there is no morphologically discrete ovuliferous scale, and there is no ontogenetic fusion of discrete separate structures. The vascular system of the scale complex develops after the ovules are initiated and forms a single series of vascular bundles with inversely oriented xylem and phloem; this and other histological changes are the result of the activity of the intercalary growth process and do not relate to primary structures. The results are discussed in relation to existing knowledge of cone development in Cupressaceae, in which axially borne ovules are common.     

Biochemical Systematics of Gymnosperms (4)—Seed Protein Peptides and Needle Peroxidases of Taxodiaceae

An analysis of seed protein peptides by using SDS polyacrylamide gel electrophoresis, indicates that all members in Taxodiaxeae share three peptides with molecular weights of 24, 32 and 10 kilodaltons. Generally, seeds of this family contain 2 to 4 major peptides, and their molecular weights are distributed in two regions: 24-26 K and 29-36 K respectively. The quantity of these major peptides is near or even more than 50% of seed total protein content. This kind of protein profiles is similar to that of Taxaceae and Cephalotaxaceae rather than to that of Pinaceae. On the other hand, a considerable difference was observed in peptides with higher molecular weights. Among different species within a genus, no difference has been found both in their seed protein profiles and in their needle peroxidases. Based on the data cited here, it seems that the relationships among most genera of Taxodiaceae are not so closed as the rela-tionship among following genera: Sequoia, Sequoiadendron and Taxodium.     

A taxodiaceous seed cone from the Triassic of Antarctica

A silicified seed cone is described from the lower Middle Triassic of A silicified seed cone is described from the lower Middle Triassic of Antarctica. The cone measures up to 3.4 cm long and 1.4 cm wide, and consists of helically arranged cone scales attached to a eustelic axis. Bract and ovuliferous scale are approximately of equal length and fused at the base. The bract is entire and vascularized by a single trace. The ovuliferous scale contains five distal lobes, each vascularized by a terete strand that divides to form a smaller trace to each of the five inverted ovules. Ovules are small and flattened with the three-parted integument attenuated into oppositely positioned lateral wings. The Triassic specimens are compared with both extant and fossil conifer seed cones and believed to have their closest affinities within the Taxodiaceae.     

CONE AND OVULE DEVELOPMENT IN SCIADOPITYS (TAXODIACEAE-CONIFERALES)

Ontogeny of seed cones of Sciadopitys, with special reference to the ovule-supporting structure, is studied in material collected in Japan and Massachusetts. Cones are initiated as lateral or terminal structures in summer and complete the formation of most organs before winter. Bract development is well advanced before ovule-supporting structures are initiated. Continued cone development involves the formation of a narrow ridge of tissue in the axil of each fertile bract. This ridge develops a series of nine (but up to 12) apical lobes in centrifugal order, each of which is the primordium of a future tooth on the ovuliferous scale. Ovules are initiated as outgrowths of the adaxial surface of each lobe so that there is a one-to-one ratio between lobes and ovules. Intercalary extension of the ovuliferous scale itself (distally) and the common base of the bract and ovuliferous scale (proximally) greatly extends the complex. The ovuliferous scale eventually exceeds the subtending bract and its apex becomes recurved. Bracts each have a single trace, but each ovuliferous scale has a pair of traces that proliferate distally to irrigate ovule and scale lobe. Intercalary growth results in recurvature of the ovule trace. The organization of the cone is directly comparable with certain Permian fossils. Sciadopitys also seems unique within the Taxodiaceae in its centrifugal development of the ovule-supporting complex.     

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。     

北美香柏雌球果的发育

用扫描电镜(SEM)观察了北美香柏 Thuja occidentalis 雌球果的发育过程。在北京,北美香柏的雌球果是在八月初由营养芽转变而来,雌球果一般有4~6对苞片,中间2~3对可育,每一苞片腋部着生两枚胚珠,在可育苞片腋部最先观察到一扁平的隆起,并在其上分化出两个胚珠原基,接着分化出珠被和珠心,最后形成扁平而两侧对称的胚珠。在北美香柏雌球果发育过程中,约一半的雌球果在2~3对可育苞片中位于下面的1~2对的腋部产生3个胚珠原基,中间一个较小,并在以后的发育中逐渐退化。由此推测北美香柏的雌球果可能是由祖先类群中每一苞片具多于2个胚珠的雌球果演化而来。在光镜下对雌球果维管系统的观察发现,传粉前幼小雌球果的苞片内仅有一束维管束,传粉后随着苞片基部的居间生长,有4—8束维管束在苞片内形成,但是新发育的维管束木质部和韧皮部相对位置与正常叶性器官一致,这与在以往报道的柏科植物成熟雌球果的苞片中均有反向维管束的发育不同。北美香柏雌球果早期发育和维管束分析结果支持傅德志和杨亲二提出的解释裸子植物生殖器官形态演化的“苞鳞-种鳞复合体”理论。关键词北美香柏;雌球果的发育;胚珠分化;SEM