松科植物属间C值和种子千粒重与进化关系初探

在检索C值数据库和种子数据库的基础上,对松科具有完整C值和种子千粒重等数据信息的5个属108种植物进行统计分析,并利用线性回归模型分析了二者之间相关性。结果显示,松科植物C值从9.5~36 pg呈正态分布,种子千粒重从1.2~1 269 g呈偏态分布,种间千粒重相差悬殊,高达1058倍。ANOVA分析表明,松属C值平均为25.79 pg,显著高于其它4个属;落叶松属C值最低（12.53 pg),显著低于云杉属（18.44 pg）,而云杉属、冷杉属和雪松属C值间却无显著性差异。结合前人研究结果,从C值的角度分析,松科5属间的演化顺序为：松属→云杉属→雪松属→冷杉属→落叶松属。松科5属间种子千粒重均值差异显著,松属种子均值最大（123.7 g）,其次为雪松属（84.5 g）和冷杉属（26.4 g）,最后为落叶松属（5.5 g)和云杉属(3.9 g)。除松属外,其它4个属ANOVA分析,雪松属种子千粒重显著高于云杉属、冷杉属和落叶松属（P<0.01),而云杉属和落叶松属种子千粒重间无显著差异(P>0.05)。对数回归分析和线性相关性分析表明,松科108种植物的C值和种子千粒重回归方程为：y=3.145x-7.248,决定系数（R²）为0.346,存在显著正相关性（P<0.000 1）,线性相关系数(γ)高达58.8%,这说明松科植物随着C值增大,其种子千粒重呈明显增加趋势。综上所述,松科属间种子千粒重差异显著,且随着属间进化程度的提高,C值和种子千粒重均呈明显下降趋势,其具体机理尚有待于结合其它功能性状做深入分析。     

SEM study of the inner periclinal surface of leaf cuticles in the family Pinaceae

Characters of the internal cuticle surface of 82 species of Pinaceae (ranging over nine genera) have been investigated using both light and SEM. Considerable variation is demonstrated in details of the intercellular flanges and in the fine sculpturing of the periclinal surface. Nine different sculpturing patterns are recognized. There is systematic value in this character; species within genera can usually be distinguished on the basis of cuticle structure, except in Cedrus and Larix .     

Cytogenetic and molecular characterization of the Abies alba genome and its relationship with other members of the Pinaceae

Genome size, karyotype structure, heterochromatin distribution, position and number of ribosomal genes, as well as the ITS2 sequence of the internal transcribed spacer (ITS) were analysed in silver fir (Abies alba Mill.). The analysis also included characterization of the Arabidopsis-type of telomeric repeats in silver fir and in related species. The results were compared with results from other species of the Pinaceae, to evaluate phylogeny and chromosomal and molecular evolution in the Pinaceae. Integrated chromosomal data provided insights into chromosome and karyotype evolution in the Pinaceae. The evolutionary trend for GC-rich heterochromatic blocks seems to involve loss of blocks that are not associated with rDNA. Similarly, numerous large blocks of interstitial plant telomeric repeats that are typical for all analysed species of the genus Pinus were not observed in the evolutionarily younger genera, such as Abies, Picea and Larix. On the contrary, the majority of telomeric sequences in these three genera appeared confined to the chromosome ends. We confirmed the current position of Abies and Tsuga in subfamily Abietoideae and the position of Pinus in the subfamily Pinoideae based on ITS2 sequences. Pseudotsuga is placed together with Larix into the subfamily Laricoideae. We conclude that the current position of the genus Picea in the subfamily Abietoideae should be reconsidered and, possibly, the genus Picea should be reclassified as a separate subfamily, Piceoideae, as recently proposed.     

The course of resin canals in the shoots of conifers

Resin canal patterns in the Pinaceae differ significantly from those in other conifers. In this paper, Pinus type, Picea type and Larix type are defined.Pinus has the Pinus type pattern, in spite of the possession of a system of peripheral canals in some members. The patterns inAbies, Picea, Cedrus, Keteleeria andPseudotsuga are varied. Both Pinus and Picea types are usually observed, although many other types are also seen in these five genera.Larix andPseudolarix have the Larix type pattern, andTsuga has a modified Torreya type pattern. From the data obtained in the present series of studies (Suzuki, 1979a, b), two main evolutional trends of resin canal patterns are suggested. In the Pinaceae, the patterns probably evolved from the complicated type to the simple type. In the other families, they evolved from the simple type to the complicated type. Moreover, it is to be considered likely that the patterns have evolved independently in each family. Patterns of resin canals are very similar to the primary vascular patterns. The similarity is probably not fundamental, but superficial, because most of the conifer species, of which resin canal patterns are varied, have but a single type of the primary vascular system.     

松科系统发育的分子生物学证据

运用PCR方法分别从松科8属、9种植物中扩增出一长约2550bp的cpDNA片段,这一片段包括rbcL、trnR、部分accD及基因间的非编码区(相对于黑松cpDNA中的同源片段而言)。运用18种限制性内切酶对这一cpDNA片段进行酶切分析,共获得86个酶切位点,其中54个为变异位点。运用PAUP(version 3．1．1)和Mega(version 1．01)软件对数据进行分析,结果Wagner简约树和Neighbor-Joining树反映出的松科系统发育关系基本一致：银杉属、松属、黄杉属和落叶松属形成一个单系群,且银杉属与松属的亲缘关系更近于与另外二属的关系,但这一结果未得到Bootstrap分析的较强支持;落叶松属近缘于黄杉属;冷杉属近缘于油杉属。此外,松科中的冷杉亚科和落叶松亚科均不是单系类群,将松科划分为冷杉亚科、落叶松亚科和松亚科三个亚科的系统(郑万钧,傅立国,1978)是不自然的。     

Areography of the Gymnosperms of China (1) Distribution of the Pinaceae of China

Nine of 10 genera and 119 of approximately 240 species of the Pinaceae occur in China, including 67 endemic species and two endemic genera. In this paper, the distributional maps of all the genera of the Pinaceae are presented (fig. 1-8). The horizontal and vertical distributions of species in each genus are discussed. The analysis of the distribution patterns of the genera indicates that some genera, such as Keteleeria, Tsuga, Pseudotsuga, Cathaya and Pseudolarix, are restricted to the area south of the Qinling Mountains and the Huaihe River, and the others, i. e. Picea, Abies, Larix and Pinus, extend northward to northeastern China. However, all of the genera except Keteleeria and Pinus are not found in very dry areas and tropical mountainous regions of China. The monotypic genera, Cathaya and Pseudolarix, are distributed in eastern and central China. The genus Keteleeria consists of 10 species, 7 of which are concentrated in southern Guizhou, northern Guangxi, southwestern Hunan and easternmost Yunnan. The distribution of the remaining 6 genera shows the maximum concentration in western Sichuan and northwestern Yunnan. (Figs. 2-8). Furthermore, more than third of species of the Pinaceae (37.8%) are also concentrated in western Sichuan and northwestern Yunnan. where a great variety of habitats and different topographic features occur. It is apparent that to conduct our systematic and evolutionary studies on this family in these region is especially needed. The relations between the areal size and the tolerance of species are discussed. The distributions of macrofossils and microfossils of the genera of the Pinaceae ia China are given, and it has been proved that areas of most genera of the family were considerably larger in the past. than at present.     

Comparative wood anatomy in Abietoideae (Pinaceae)

This study, which includes 51 species and six genera of subfamily Abietoideae (Pinaceae), assesses the systematic significance of the wood structure in this group. In particular, the presence of normal and traumatic resin canals, the ray structure and the axial parenchyma constitute phylogenetically informative features. Comparative wood anatomy of Abietoideae clearly supports the monophyly of the genera Abies–Cedrus–Keteleeria–Nothotsuga–Pseudolarix–Tsuga, all of which have axial parenchyma with nodular transverse end walls in the regions of growth ring boundaries, crystals in the ray parenchyma and pitted horizontal and nodular end walls of ray parenchyma cells. Axial resin canals support a subdivision of the subfamily into two groups: Abies, Cedrus, Pseudolarix and Tsuga, without axial resin canals, and Keteleeria and Nothotsuga, with axial resin canals and a specific arrangement of traumatic axial resin canals. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 184–196.     

松科系统学研究(Ⅰ)

松科的系统发育研究历来已久,但由于性状进化的不同步性和属间进化缺少中间环节,致使科的分类系统和各属之间的演化关系至今未得到令人满意的解决。本文在钻研已有文献的基础上,筛选出松科属间综合分类性状36个,有序进化性状20个,借助计算机,以属为分类单位,进行表征分析研究确定松科分类系统,结果将松科分为三个亚科:松亚科Subfam.Pinoideae:仅Pinus Linn;落叶松亚科Subfam.Laricoideae Melchior & Werderm.emend.N.Li ex L.K.Fu:含四属:Cathaya Chun & Kuang, Larix Mill., Picea A.Dietr., Pseudotsuga Carr.;冷杉亚科Subfam.Abietideae Pilger, emend.Frankis:含五属:Abies Mill., Cedrus Trew, Keteleeria Carr., Pseudolarix Gordon, Tsuga Carr.。运用分支分析探讨各属的演化关系,并综合表征和分支分析结果,结合古植物学资料,构思和绘出了以地质年代为纵作标的松科系统发育树。     

Comparative anatomy of resin ducts of the Pinaceae

 Resin ducts are common in the Pinaceae. The comparative anatomy of stems and leaves of 50 species and two varieties from ten genera has been investigated. The structure and distribution of resin ducts differ among genera. Resin ducts occur in foliage leaves of ten genera of Pinaceae. Cortical resin ducts are absent in the stems of Pseudolarix and Larix. Resin ducts only occur in the secondary xylem of stems of Pinus, Picea, Cathaya, Larix, Pseudotsuga and some Keteleeria species. All of the epithelial and sheath cells are alive and thin-walled in the resin ducts of stem cortex and mesophyll. Except for Pinus the epithelial cells of resin ducts in the secondary xylem of stems have thick, lignified walls. Comparative study shows there are obvious differences in the resin ducts of different genera; apparent differences do not exist, however, in the resin ducts of different species of the same genus. According to the structure and distribution of the resin ducts in ten genera of Pinaceae, a synoptical arrangement of the genera is given and generic relationships of the Pinaceae are discussed. Received: 12 September 1995 / Accepted: 14 March 1996     

松科4属植物茎初生结构比较研究

研究比较观察了松科云杉属的云杉（Ｐｉｃｅａ ａｓｐｅｒａｔａ）、油杉属的油杉（Ｋｅｔｅｌｅｅｒｉａ ｆｏｒｔｕｎｅｉ）、雪松属的雪松（Ｃｅｄｒｕｓ ｄｅｏｄｅｒａ）、松属的海南五针松（Ｐｉｎｕｓ ｆｅｎｚｅｌｉａｎａ）和大明松（Ｐ．ｔａｉｗａｎｅｎｓｉｓ Ｈａｙａｔａ ｖａｒ．ｄａｍｉｎｇｓｈａｎｅｎｓｉｓ）５个种的幼茎初生结构。结果表明,皮下层的细胞层数、皮层细胞的组成,树脂道的分布,鞘细胞的排     

Length Variation of the Nuclear Ribosomal DNA Internal Transcribed Spacer in the Genus Abies,with Reference to Its Systematic Utility in Pinaceae

The internal transcribed spacer (1TS) region (1TS1, ITS2 and 5.8S rDNA) of the nuclear ribosomal DNA (nrDNA) was amplified via PCR in 28 taxa of Abies Mill. The amplified fragments showed length polymorphism among species, with species from Central America and two species from North America having a length of approximately 2 500 base pairs (bp) and the remaining taxa having a length of approximately 1 700 bp based on 100 bp and 1 kb ladder standard markers. The complete sequencing of ITS of Abies bracteata showed that the shorter type is 1 697 bp (1TS1 is 1 296 bp, 5.8S + 1TS2 is 401 bp). For the longer one, the partial rrs1 and complete 5.8S + ITS2 sequencing revealed that thelength of 5.8S + ITS2 is the same as that of the shorter type. The length difference of ITS in Abies is mainly due to the length difference in the ITS1 region, a result similar to the previous findings in other genera of Pinaceae. Variation in ITS length seems well correlated with morphological and geographic characters in Abies, suggesting that the length variation may be a phylogenetically informative character within the genus, long ITS was also found in other genera of Pinaceae in the previous studies. The long length of ITS in the family makes the sequencing of the region and subsequent alignment of sequences among species or genera more difficult than in taxa with short ITS, such as angiosperms. Although the length variation of ITS in the genus Abies is significant, the homogenous of ITS sequence between the longer one and the shorter one is obvious if the insertion in the longer ITS is ignored.     

Fundamental features of the distribution of Coniferae in Sichuan

The abundance of Coniferae in Szechuan Province is a well-known fact, especially of the Order Pinales. There are so far 19 genera 68 species and varieties belonging to the Pinaceae, Taxodiaceae and Cupressaceae. This paper dealing with the fundamental features in the distribution of these plant-groups may be not only of scientific interest but also of great importance to forestry. According to an analysis of the influence of historic, geographic and ecological conditions on the distribution of plants, it is shown that the differantiation of geographical distribution of genera and species in these 3 familias between the eastern and the western areas is very evident. There are (Tab. 1) 18 genera and 27 species in the eastern area, including 6 monotypic genera, they are Cathaya, Pseudolarix, Metosequoia, Glyptostrobus, Fokienia and Platycladus. As the great concentration of the monotypic and paleo-endemic genera is not found elsewhere, this area therefore distinctly indicates the relic nature. It seems most probably that they came originally into existence in subtropical and warm-temperate climates of middle lalitudes in the northern hemisphere, and most of them are in the lands bordering the Pacific Ocean. On the other hand, there are 13 genera and 56 species in the western area with only one monotypic genus (Platycladus) present, and most genera are polytypic with wide ranges. The number of species in these genera here is larger than in any other areas, for example, Abies and Picea are represented 12 species each (Fig. 1-4). There are 10 species in Sabina (Fig. 5). It might be considered that each genus has developed into a great centre, In addition, it has been discussed in detail regarding the vicarious distributions in geography, verticality, and ecology of ralated species in Abies, Picea, Tsuga, Keteleeria, Larix, Pinus and Cupressus (Fig. 3-5), as well as the re lationship of these species. As shown above, these plants are also, without doubt, of very old origin. Moreover, they are most likely very much closely related with Tethys in the past, but the majority species apparently have arisen in relalively recent times. They are due to the effect of mountain-making and glaciation, principally in consequence of the cooling process of the climatic vicissitude.     

中国松科花粉形态研究

本文研究了我国松科(Pinaceae)9属61种的花粉形态, 其中有6种进行扫描电子显微镜观察和照相, 4种进行透射电子显微镜观察和照相, 每属都有代表种的绘图, 对该科花粉形态类型、科下的分类及演化问题进行讨论, 为本科的亚科、属、种的划分提供孢粉学方面的参考依据。     

Two highly divergent 5S rDNA unit size classes occur in composite tandem array in European larch (Larix decidua Mill.) and Japanese larch (Larix kaempferi (Lamb.) Carr.).

The 5S ribosomal DNA unit structure and organization have been investigated in Larix decidua and Larix kaempferi using selective amplification of gene and spacer, sequence analysis and homologous probe hybridization. Two highly divergent unit size classes of approximately 650 and 870 bp were detected in both species. Sequence analysis in Larix decidua revealed that length variations occur in the middle spacer region and are the result of duplications (in the long spacers) and considerable sequence heterogeneity. Conversely, the transcribed region is of uniform length (120 bp), and the nucleotide sequence of one Larix decidua clone is similar to that reported for other gymnosperms. Sequence comparison of the larch spacers with two other Pinaceae species (Pinus radiata and Picea glauca) showed that the 5' and 3' regions flanking the gene (40 and 60 bp, respectively) are quite conserved, suggesting a regulatory role. Moreover, a small element of about 70 bp located in the middle spacer region was found to be common to the larch long units and the six Pinus radiata spacer clones previously sequenced (64% sequence identity). The short and long unit size classes are mainly organized in composite tandem array(s) with evidence of extensive zones of strict alternation in both species. Mechanisms underlying this unusual association of divergent units in larch 5S rDNA arrays are discussed.     

A world revision of Strobilomyia gen.n.: the anthomyiid seed pests of conifers (Diptera: Anthomyiidae)

Abstract. The world species of anthomyiid cone and seed pests are revised. A new genus, Strobilomyia gen.n., is erected for the eighteen species, including seven new to science, presently recognized. Author's views on homologies in the male and female terminalia of Anthomyiidae and other Diptera Cyclorrhapha are amplified as background for the terminology employed. The monophyly and systematic position of Strobilomyia are discussed with reassessions of related genera: Lasiomma Stein, Chirosia Rondani, Egle Robineau-Desvoidy and Acklandia Hennig. Zoogeographic aspects, especially vicariance across the Beringia, are analysed from a partial hypothesis of phylogenetic relationships among the species of Strobilomyia in conjunction with paleo-environmental data. New generic synonyms are: Macrophorbia Malloch, Crinurina Karl, Si-nohylemya Hsue (all =Lasiomma); Acrostilpna Ringdahl, Shakshainia Suwa, Meliniella Suwa (all =Chirosia). The species recognized in Strobilomyia (with geographic range and host trees in parentheses) are: anthracina (Czerny) (Palearctic, Picea); neanthracina sp.n. (Nearctic, Picea and ITsuga); appalachensis sp.n. (E. Nearctic); melania (Ackland) (Europe, Larix); sibirica sp.n. (N.E. Europe, Siberia, Larix); viaria (Huckett) (E. Palearctic, Nearctic, Larix); infrequens (Ackland) (Palearctic, Larix); luteoforceps (Fan & Fang) (E. Palearctic, Larix); baicalensis (Elberg) (E. Palearctic, Larix); laricicola (Karl) (Palearctic, Larix); laricis sp.n. (Nearctic, Larix); macalpinei sp.n. (W. Canada, Larix); carbonaria (Ringdahl) (C. Europe, E. Nearctic, Abies); svenssoni sp.n. (N. Europe, Mongolia); suwai sp.n. (Japan); todocola (Suwa) (Japan, Abies); abietis (Huckett) (Nearctic, Abies); and oriens (Suwa) (E. Palearctic, Abies).     

Genetic diversity and structure of natural and managed populations of Cedrus atlantica (Pinaceae) assessed using random amplified polymorphic DNA

Cedrus atlantica (Pinaceae) is a large and exceptionally long-lived conifer native to the Rif and Atlas Mountains of North Africa. To assess levels and patterns of genetic diversity of this species, samples were obtained throughout the natural range in Morocco and from a forest plantation in Arbúcies, Girona (Spain) and analyzed using RAPD markers. Within-population genetic diversity was high and comparable to that revealed by isozymes. Managed populations harbored levels of genetic variation similar to those found in their natural counterparts. Genotypic analyses of molecular variance (AMOVA) found that most variation was within populations, but significant differentiation was also found between populations, particularly in Morocco. Bayesian estimates of F(ST) corroborated the AMOVA partitioning and provided evidence for population differentiation in C. atlantica. Both distance- and Bayesian-based clustering methods revealed that Moroccan populations comprise two genetically distinct groups. Within each group, estimates of population differentiation were close to those previously reported in other gymnosperms. These results are interpreted in the context of the postglacial history of the species and human impact. The high degree of among-group differentiation recorded here highlights the need for additional conservation measures for some Moroccan populations of C. atlantica.     

Serine proteinase inhibitors in seeds of Cycas siamensis and other gymnosperms

Seeds of 32 species selected from two of the four major groups of gymnosperms, the ancient Cycadales and the economically important Coniferales, were analysed for inhibitors (I) of the serine proteinases trypsin (T), chymotrypsin (C), subtilisin (S) and elastase (E) using isoelectric focusing (IEF) combined with gelatin replicas. Subtilisin inhibitors were detected in 17 species, being particularly active in the Cycadales. Several species of the genera Cephalotaxus, Pseudotsuga and Cycas contained inhibitors active against elastase while strong CSTIs and CSIs were also present in Cycas pectinata and C. siamensis. No inhibitors were detected in seeds of Chamaecyparis, Thuja, Abies, Larix, Picea and Pinus spp. Serine proteinase inhibitors were purified from seeds of C. siamensis by affinity chromatography using trypsin and chymotrypsin, IEF and SDS-PAGE. Several CSTI components with M_r ranging from 4000 to 18,000 were partially sequenced using Edman degradation and mass spectrometry. Most of the sequences were similar to a hypothetical protein encoded by an mRNA from sporophylls of C. rumphii which in turn was similar to Kunitz-type proteinase inhibitors from flowering plants. Analysis of expressed sequence tag (EST) databases confirmed the presence of mRNAs encoding Kunitz-type inhibitors in the Cycadales and Coniferales and also demonstrated their presence in a third major group of gymnosperms, the Ginkgoales. This is the first report of Kunitz-type serine proteinase inhibitors from plants other than Angiosperms.     

Genetic variability of Siberian fir (Abies sibirica Ledeb.) inferred from AFLP markers

Genetic variability of AFLP markers was studied in 20 populations of Siberian fir (Abies sibirica (Pinaceae) and in two populations of Far-Eastern Manchurian fir A. nephrolepis and Sakhalin fir A. sachalinensis each. Four pairs of selective primers were used. In total, 168 samples from three fir species were genotyped for 117 polymorphic loci. According to the AMOVA results, the variability proportion characterizing the differences between three Abies species was several times higher (F(CT) = 0.53) than that accounting for among-population differences within the species (F(SC) = 0.125). Differentiation of the A. sibirica populations based on AFLP markers exceeded 14% (F(ST) = 0.141). Significant correlation between the genetic distances calculated from the AFLP data and the geographic distances between populations was found. The results of AFLP variability analysis supported and supplemented the conclusions inferred previously from allozyme and cpSSR data: several genetically similar geographic groups of Siberian fir were identified. These groups differ both in allele frequencies and in the levels of genetic variation.     

Distribution and hosts of Callidiellum rufipenne (Coleoptera: Cerambycidae), an asian cedar borer established in the eastern United States

The distribution and hosts of the exotic cedar-boring beetle, Callidiellum rufipenne (Motschulsky) (Coleoptera: Cerambycidae), were determined in five northeastern U.S. states by capturing adults on cedar trap logs and by rearing adults from various conifers. This beetle was detected in the coastal states of Massachusetts, Rhode Island, Connecticut, New York, and New Jersey. In these states, adults emerged from the live or dead wood of four genera and eight species of Cupressaceae; species of Pinaceae were not hosts. Through its entire range, C. rufipenne is reported to infest at least 14 species of Cupressaceae, four species of Pinaceae, and one species of Taxaceae; but, records of Pinaceae and possibly Taxaceae are suspect. Based on the number of adults that emerged from coniferous poles in a five-way choice test in the field, the infestation level was significantly greater in Chamaecyparis thyoides (L.) Britton, Sterns, and Poggenburg and Juniperus virginiana L. than in Pinus rigida Miller, Pinus strobus L., and Tsuga canadensis (L.) Carribre (last three species uninfested). In a second test of host preference in the wild, beetles infested four cupressaceous species, but not Abies balsamea (L.) Miller, Picea rubens Sargent, Pinus rigida, P. strobus, and Ts. canadensis in the Pinaceae. Infestation level was highest in Ch. thyoides, followed in decreasing order by Juniperus communis L., Thuja occidentalis L., and J. virginiana. In a comparison of live and dead J. virginiana, beetles developed to adults only in dead trees (36 beetles per tree). When trunk sections of Th. occidentalis with and without bark were offered to females in cages, beetles of the next generation emerged exclusively from wood with bark. In the Northeast, only species of Cupressaceae apparently are suitable hosts for C. rufipenne. Infestation of these species may be prevented or reduced by proper care of live plants and by debarking trees after harvesting.