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 .     

Fossil History and Modern Distribution of the Genus Abies (Pinaceae)

The plants of the genus Abies are dominant and key species in dark conifer forest in the Northern Hemisphere. There are 52 species, 1 subspecies and 12 varities of genus Abies in the world. The history and modern distribution ofAbies were discussed at present paper. The genus has 3 modern distributional centers: South Europe, North America and East Asia. These areas are also rich in fossil records. The vertical distribution regions of Abies are from sea level to 4 700m, concentrated in 1 000 - 2 000 m (15 species ) and 2 500 - 4 000 m ( 13 species ). In China, the genus distributes in 20 provinces, especially abundant in the Hengduan Mountians. Meanwhile endemic and relic phenomea are obvious in this genus. There are 7 relical species with both limited individuals and limited distributed regions. Based on the fossil records and the newest phylogenetic data, the following hypothesis was proposed: Abies originated from the mid- and high altitude of the Northern Hemisphere in the Middle Cretaceous and it was dispersed forward to the south area in the Eocene due to global climate cooler down. The distribution of Abies was deeply impacted by geological events such as upleft of Himalaya, Alps, Rocky Mountains, the occurrence of Aisan Monsoon as well as Quaternary glaciers. Finally the currentdistribution pattern appeared at the Quaternary. The genus Abies has similar fossil history and modern distribution pattern with Cathaya and Pseudolarix.     

Systematic wood anatomy of the tribe Guettardeae (Rubiaceae)

Systematic wood anatomy of the tribe Guettardeae (Rubiaceae). The wood anatomy of nearly all genera of the Guettardeae (Rubiaceae, Guettardoideae) has been examined, and in this respect the tribe is heterogeneous. Suggestions are made for a delimitation of the tribe. Guettarda, Bobea, Antirhea, Malanea and Chomelia Jacq. are sufficiently similar in their wood anatomical characters to warrant retention in the same tribe. Machaonia, Timonius and Dichilanthe are anomalous. Suggestions are given for taxonomic revisions of some genera based on their wood anatomy.     

Isolation and characterization of polymorphic microsatellite markers in the endangered Nothotsuga longibracteata (Pinaceae)

Twelve polymorphic microsatellite loci were isolated and characterized from an AC‐enriched genomic library of Nothotsuga longibracteata. The average allele number of these microsatellites was 8.3 per locus, ranging from two to 13. The ranges of observed and expected heterozygosities were 0.03–0.97 and 0.09–0.88, respectively. These polymorphic markers provide useful tools for the study of evolutionary history and conservation genetics of N. longibracteata.     

上新世云南铁杉木化石的化学成分

为了研究在褐煤中发现的上新世云南铁杉(Pliocene Tsuga dumosa)木化石的化学成分,采用气相色谱-质谱联用技术和柱层析方法分别检测了22个挥发性成分和分离鉴定了9个化合物(1-9),化学成分类型包括烷烃(C16-C32)、脂肪酸、甾体、三萜、邻苯二甲酸酯及酚性成分。化合物结构采用波谱方法鉴定。     

Germination as a determinant of seedling distributions among natural substrates in Picea engelmannii (Pinaceae) and Abies lasiocarpa (Pinaceae)

In Rocky Mountain (USA) subalpine forests, seedlings of Picea engelmannii (Engelmann spruce) colonize logs more frequently than seedlings of its codominant associate Abies lasiocarpa (subalpine fir). We hypothesized that spruce germinates more readily on logs than fir, perhaps because small spruce seeds are more likely to lodge in log crevices than larger fir seeds. Our objectives were to test this hypothesis and compare both species' germination among several substrates to assess germination influences on natural seedling distributions. Spruce and fir seeds were sown on field-collected logs, litter, and soil in the greenhouse and monitored for 36 d. To test the crevice hypothesis, seeds were either scattered on logs or wedged into crevices, assuming that if both species were placed in crevices, interspecific germination differences on logs would decrease. Spruce mean germination percentages were significantly greater than fir's in all substrate treatments except when seeds were wedged in log crevices. The difference in means between the two log treatments was greater for fir (68%) than spruce (21%). Spruce germinated more rapidly than fir on all substrates. We suggest that large seed size reduces fir's success in colonizing logs, and that germination and establishment factors interact to determine natural seedling distributions for these subalpine conifers.     

Vesicular endophytes in roots of the Pinaceae

Vesicles and hyphae typical of vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii, Abies lasiocarpa and Tsuga mertensiana growing in openings where herbaceous hosts of these fungi were common. Seedlings of A. lasiocarpa, Tsuga heterophylla, and T. mertensiana growing under closed forest canopies also had vesicles but at a much lower incidence than seedlings in the openings. The Pinaceae are generally assumed to be ectomycorrhizal, but Glomus-type colonizations occurred on the same seedlings as the ectomycorrhizae. The ecological significance of abundant VAM-type endophytes in otherwise ectomycorrhizal hosts deserves comprehensive study.     

Comparative wood anatomy ofCoriaria

Wood anatomy ofCoriaria was surveyed to clarify generic features on the basis of 14 species collected from various regions of the World to cover the whole range of geographic distribution and habitual variation. Wood anatomy ofcoriaria is considerably uniform, and the species share a combination of the following features: 1) pores are thin-walled, polygonal in outline and mostly in multiples; 2) vessel elements and libriform fibers are very short; 3) perforation plates are exclusively simple; 4) intervessel pits are alternate; 5) vascular tracheids are present; 6) wood parenchyma is vasicentric and sometimes confluent; 7) rays are heterogeneous and large. Its species differ in several characters, such as distinctness of growth rings, pore size, pore patterns, type and abundance of wood parenchyma, and distinctness of storied structure. Comparisons among species indicate that the species of the Northern Hemisphere show a tendency toward having semi-ring porosity, while those of the Southern and Western Hemisphere have diffuse porosity. The other infrageneric variations appear to be related to different habits of the species rather than to geographic distribution. Small trees mostly have confluent and vasicentric parenchyma composed of fusiform cells and distinctly storied tissues, while shrubs and herbs have less abundant parenchyma which is vasicentric and comprises strands of two to four cells and indistinctly storied tissues.     

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     

松科冷杉属植物的化石历史和现代分布

冷杉是北半球阴暗针叶林的优势种和建群种,现全世界共有52种1亚种12变种,在北半球形成南欧、北美和东亚三个分布中心,这三个地区也是冷杉属化石最丰富的地区。在垂直分布上,冷杉集中分布于1000～2000m（15种）和2500～4000m（13种）两个海拔地段。在中国,冷杉植物呈南北间断分布,集中分布在横断山地区。冷杉属的特有现象和孑遗分布现象都十分突出,有7个种呈孑遗分布。根据冷杉属的地史分布和现代分布的研究并结合最新的系统演化资料,本文推测冷杉属于白垩世中期起源于北半球的中高纬度地区,始新世以后,随着全球气候的变冷,逐步向南迁移,由于喜马拉雅山脉、阿尔卑斯山、落基山脉抬升及东亚季风气候的出现以及第四纪冰期的影响而形成了现代间断的分布格局。冷杉与银杉、金钱松等其它松科植物的形成模式十分相似。     

Genetic variation in Tertiary relics: The case of eastern‐Mediterranean Abies (Pinaceae)

The eastern‐Mediterranean Abies taxa, which include both widely distributed species and taxa with minuscule ranges, represent a good model to study the impacts of range size and fragmentation on the levels of genetic diversity and differentiation. To assess the patterns of genetic diversity and phylogenetic relationships among eastern‐Mediterranean Abies taxa, genetic variation was assessed by eight nuclear microsatellite loci in 52 populations of Abies taxa with a focus on those distributed in Turkey and the Caucasus. Both at the population and the taxon level, the subspecies or regional populations of Abies nordmanniana s.l. exhibited generally higher allelic richness, private allelic richness, and expected heterozygosity compared with Abies cilicica s.l. Results of both the Structure analysis and distance‐based approaches showed a strong differentiation of the two A. cilicica subspecies from the rest as well as from each other, whereas the subspecies of A. nordmanniana were distinct but less differentiated. ABC simulations were run for a set of scenarios of phylogeny and past demographic changes. For A. ×olcayana, the simulation gave a poor support for the hypothesis of being a taxon resulting from a past hybridization, the same is true for Abies equi‐trojani: both they represent evolutionary branches of Abies bornmuelleriana.     

SEM Observation on the Structure of Cuticles on Leaf Inner Surface of Abies (Pinaceae) and Its Significance in Systematics

Comparative investigation of the inner surface of the needle cuticle of 36 species and 2 varieties of Abies under SEM has revealed that the characteristics of the intercellular flanges are rather distinct and four types can be distinguished: (1) Straight and developed single flange. This type is only represented by Abies bracteata D. Don. Morphologically, this species is also quite unique in the genus Abies and was once treated as a subgenus by Franco and Liu. Its special structure of the leaf cuticle observed here seems to support their treatment. (2) Double flanges. This type was first discovered in a leaf fossil of Abies from England. In modern plants of Abies, it is found only in the species from Central America. (3) Undeveloped single flange. This type is represented by a small group of Abies from the west and east coastal area of the Pacific Ocean. (4) Undulate and developed single flange. This type is represented by most of the species of Abies, including all the species in Europe and most species in Asia and North America. The flange types mentioned above seem to have some relationships with the geographical distribution of the species in the genus Abies, and their occurrence might have not been completely influenced by the habitats, hence the features of the intercellular flanges may provide good evidence for the subgeneric division of Abies. Based on our results and those from the previously published literature about the infrageneric treatments of Abies and the distribution of the fossils, we consider that western North America might be the diversity center of modern Abies. Florin once pointed out that the characters of the leaf cuticle in gymnosperms are of great significance for the generic andinfrageneric division. This viewpoint is strongly supported by our study on modern Abies.     

Phylogeny and biogeography of Cedrus (Pinaceae) inferred from sequences of seven paternal chloroplast and maternal mitochondrial DNA regions

BACKGROUND AND AIMS: Cedrus (true cedars) is a very important horticultural plant group. It has a disjunct distribution in the Mediterranean region and western Himalaya. Its evolution and biogeography are of great interest to botanists. This study aims to investigate the phylogeny and biogeography of Cedrus based on sequence analyses of seven cytoplasmic DNA fragments. METHODS: The methods used were PCR amplification and sequencing of seven paternal cpDNA and maternal mtDNA fragments, parsimony and maximum likelihood analyses of the DNA dataset, and molecular clock estimate of divergence times of Cedrus species. KEY RESULTS: Phylogenies of Cedrus constructed from cpDNA, mtDNA and the combined cp- and mt-DNA dataset are identical in topology. It was found that the Himalayan cedar C. deodara diverged first, and then the North African species C. atlantica separated from the common ancestor of C. libani and C. brevifolia, two species from the eastern Mediterranean area. Molecular clock estimates suggest that the divergence between C. atlantica and the eastern Mediterranean clade at 23.49 +/- 3.55 to 18.81 +/- 1.25 Myr and the split between C. libani and C. brevifolia at 7.83 +/- 2.79 to 6.56 +/- 1.20 Myr. CONCLUSIONS: The results, combined with palaeogeographical and palaeoecological information, indicate that Cedrus could have an origin in the high latitude area of Eurasia, and its present distribution might result from vicariance of southerly migrated populations during climatic oscillations in the Tertiary and further fragmentation and dispersal of these populations. It is very likely that Cedrus migrated into North Africa in the very late Tertiary, while its arrival in the Himalayas would not have been before the Miocene, after which the phased or fast uplift of the Tibetan plateau happened.     

Comparative anatomy of the heart of mammals

Hearts from 20 mammalian orders were examined according to a fixed protocol to determine if the gross differences of shape and internal anatomy could be explained. Three generalizations can be made: firstly differences are most closely related to taxonomic position, secondly some features, such as the prominence of the conus of the right ventricle in animals that leap, dig or sprint regularly as part of their defence mechanism and a long, narrow left ventricle in endurance performers may be more closely related to activity than ancestry, and lastly some characteristics may be pleiotropic effects of thoracic morphogenesis.     

Genetic discontinuity revealed by chloroplast microsatellites in eastern North American Abies (Pinaceae)

Development of conservation strategies for Fraser fir (Abies fraseri) in the southern Appalachian Mountains depends in part on recognition of the extent to which Fraser fir is genetically distinct from the closely related balsam (A. balsamea) and intermediate (A. balsamea var. phanerolepis) fir. These sibling species have exhibited intergrading, clinal variation in morphological, chemical, and genetic characteristics in prior research. Chloroplast microsatellite markers were polymerase chain reaction amplified from genomic DNA samples of 78 individuals representing the geographic ranges of Fraser, balsam, and intermediate fir. Gene diversity levels at two loci ranged among taxa from 0.65 to 0.84. Allele frequencies demonstrated significant differentiation among taxa, with R(ST) values of 0.36 and 0.10. Haplotype diversity and D(SH) were highest for balsam fir and lowest for intermediate fir. A haplotype network analysis based on allele size distribution for the two loci revealed two distinct clusters of haplotypes and population-specific haplotypes. Ninety-two percent of the haplotypes in one cluster were from balsam fir and intermediate fir, and 84% of the haplotypes in the other cluster were from Fraser fir and intermediate fir. The genetic differentiation of chloroplast DNA markers provides justification for the recognition of Fraser fir as a distinct Management Unit (MU) for conservation purposes, regardless of its taxonomic classification.     

Phenylpropanoids in mycorrhizas of the Pinaceae

Tissue-specific accumulation of phenylpropanoids was studied in mycorrhizas of the conifers, silver fir (Abies alba Mill.), Norway spruce [Picea abies (L.) Karst.], white pine (Pinus strobus L.), Scots pine (Pinus silvestris L.), and Douglas fir [Pseudotsuga menziesii (Mirbel) Franco], using high-performance liquid chromatography and histochemical methods. The compounds identified were soluble flavanols (catechin and epicatechin), proanthocyanidins (mainly dimeric catechins and/or epicatechins), stilbene glucosides (astringin and isorhapontin), one dihydroflavonol glucoside (taxifolin 3′-O-glucopyranoside), and a hydroxycinnamate derivative (unknown ferulate conjugate). In addition, a cell wall-bound hydroxycinnamate (ferulate) and a hydroxybenzaldehyde (vanillin) were analysed. Colonisation of the root by the fungal symbiont correlated with the distribution pattern of the above phenylpropanoids in mycorrhizas suggesting that these compounds play an essential role in restricting fungal growth. The levels of flavanols and cell wall-bound ferulate within the cortex were high in the apical part and decreased to the proximal side of the mycorrhizas. In both Douglas fir and silver fir, which allowed separation of inner and outer parts of the cortical tissues, a characteristic transversal distribution of these compounds was found: high levels in the inner non-colonised part of the cortex and low levels in the outer part where the Hartig net is formed. Restriction of fungal growth to the outer cortex may also be achieved by characteristic cell wall thickening of the inner cortex which exhibited flavanolic wall infusions in Douglas fir mycorrhizas. Long and short roots of conifers from natural stands showed similar distribution patterns of phenylpropanoids and cell wall thickening compared to the respective mycorrhizas. These results are discussed with respect to co-evolutionary adaptation of both symbiotic partners regarding root structure (anatomy) and root chemistry. Received: 25 May 1998 / Accepted: 6 November 1998     

广义锦葵科火绳树属4个种的枝条木材解剖及其分类学意义

研究了广义锦葵科火绳树属4个种枝条的木材解剖。火绳树属枝条为散孔至半环孔材,管孔主要为单管孔和2～3个管孔组成的径列复管孔;导管间纹孔式和射线导管间纹孔式互列、小;侵填体和螺纹加厚缺如。射线主要为单列射线,2～3列射线较多;射线细胞多为方形,射线组织主要为异型,边缘直立细胞常1行;射线组织稀为同型;鞘细胞和瓦形细胞缺如。轴向薄壁组织傍管和离管型,主要为带状。晶体丰富,主要在射线、纤维和薄壁组织中。研究的4个种可以通过枝条木材解剖特征加以区分。     

Development of nuclear microsatellite markers for the Japanese conifers Tsuga diversifolia and T. sieboldii (Pinaceae)

Nuclear microsatellite markers were developed for the two Tsuga species native to the Japanese Archipelago, Tsuga diversifolia and T. sieboldii, and a population with genetic affinities to T. diversifolia on Ulleung Island, Korea. Tsuga diversifolia and T. sieboldii are widespread dominant trees of temperate and subalpine forests in Japan but to date no genetic markers have been developed for these species. Fifteen polymorphic loci were developed and characterized, of which 14 are reliably amplified in each taxon. Across both species and the Ulleung Island population, the number of alleles per locus ranged from 3 to 26 (average = 13.93) and observed heterozygosity ranged from 0.005 to 0.935 (average = 0.535). In addition, all 15 loci were successfully amplified in a single accession of the Chinese species, T. chinensis. These markers will be useful for investigating the species’ biogeography, range‐wide genetic diversity, conservation genetic issues and potential for hybridisation.     

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