我国南部5种冷杉植物木材的比较解剖学研究

应用光学显微镜和扫描电子显微镜对分布于我国南部的5种冷杉属(Abies)植物,即资源冷杉(A．ziyuanensis)、大院冷杉(A.dayuanensis)、梵净山冷杉(A.fanjingshanensis)、元宝山冷杉(A.yuanbaoshanensis)和台湾冷杉(Akawakamii)的木材结构进行了研究。从管胞分子长度、管胞分子直径、射线高度、射线频率、纹孔类型、交叉场纹孔类型、交又场纹孔数目等方面,比较了它们木材解剖特征的差异。结果表明：大院冷杉和资源冷杉在射线频率、管胞直径、管胞长度、交叉场纹孔数目方面都很类似;元宝山冷杉在管胞直径与射线长度方面与其它冷杉有显著差异。在纹孔类型、交叉场纹孔类型方面,几种冷杉都是一致的。在交叉场纹孔数目方面,大院冷杉、资源冷杉较少,而梵净山冷杉、元宝山冷杉和台湾冷杉较多。我国南部这5种冷杉的木材解剖特征与分布在华北、东北及青藏高原上冷杉的木材非常类似。为了便于区别这5种木材,根据木材解剖特征编制了检索表。     

中国珍稀濒危冷杉的发现、现状及未来

百山祖冷杉、梵净山冷杉、元宝山冷杉和资源冷杉是中国特有的珍稀濒危植物,均被列为国家Ⅰ级重点保护植物。目前,这4种濒危冷杉在我国东南部亚热带山地呈空中岛屿状间断分布,且均分布在近山顶地区。面对全球气温的不断上升,它们缺乏足够的高海拔低温生境可供迁移,取而代之的将是生境的进一步压缩和破碎化。而横断山地区由于复杂的地形和多样化的生境而成为最合适冷杉生存的栖息地,可作为这几种冷杉的迁地保护中心。     

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。     

梵净山冷杉和元宝山冷杉的叶绿体微卫星遗传多样性分析

梵净山冷杉和元宝山冷杉是极度濒危的国家一级重点保护植物.应用叶绿体微卫星标记(cpSSR)研究它们的遗传多样性,并与同属的广布种岷江冷杉的一个种群进行比较.结果表明：3对 cpSSR 引物(Pt63718、Pt30204和Pt71936)在这3种冷杉的249个个体中共检测到21个等位基因,组成35种单倍型;它们的单倍型数和有效单倍型数分别为梵净山冷杉(No＝12,Ne＝3．92),元宝山冷杉(No＝9,Ne＝3．28),两者均低于广布种岷江冷杉(No＝14,Ne＝11．57);梵净山冷杉和元宝山冷杉的稀有单倍型较少,其频率最高的单倍型分别出现在该种群的46．0％和44．1％的个体中;梵净山冷杉(He＝0．75)和元宝山冷杉(He＝0．70)的单倍型多样性也低于岷江冷杉(He＝0．97).梵净山冷杉和元宝山冷杉的叶绿体微卫星遗传多样性水平低.     

资源冷杉与元宝山冷杉针叶精油成分的比较

为比较极小种群野生植物资源冷杉(Abies ziyuanensis)和元宝山冷杉(A. yuanbaoshanensis)针叶精油化学组成成分,该文利用水蒸气蒸馏法提取资源冷杉和元宝山冷杉针叶精油,采用GC-MS分析确定其精油化学物质成分,并基于峰面积归一化法计算各组分相对含量。结果表明:分别从资源冷杉和元宝山冷杉针叶精油中鉴定出化学物质21种和22种,其中15种为两种冷杉共有成分;烯烃类物质是资源冷杉和元宝山冷杉针叶精油中含量最高的组分,占比分别为95.94%和95.02%。两种冷杉共有成分中β-蒎烯、1-石竹烯、莰烯、α-石竹烯和异松油烯具有较大开发利用价值;非共有成分方面,资源冷杉中α-松油醇和α-蒎烯以及元宝山冷杉中α-依兰油烯和叶醇均具有较大开发利用价值。以上结果为开发利用两种冷杉针叶精油提供了理论依据。     

秦岭太白山巴山冷杉种内和种间竞争特性的研究

通过对不同方法的比较分析,提出了确定巴山冷杉(Abies fargesii)植株间竞争强度和竞争范围的新方法,并利用改进的方法研究了其种内和种间竞争强度。结果表明:随对象木胸径的增大,由于巴山冷杉种群自然稀疏过程中密度调节作用,植株距离增加,种内竞争强度降低;巴山冷杉主要分布于亚高山地段,群落内太白红杉(Larixchinensis)数量较多,胸径较大,种内与种间竞争关系顺序为:太白红杉-巴山冷杉>巴山冷杉-巴山冷杉>牛皮桦(Betula utilis)-巴山冷杉>其它树种-巴山冷杉;竞争强度和对象木胸径的关系服从幂函数关系(CI=AD-B),当巴山冷杉胸径达到25 cm以上时,种内和种间竞争强度变化较小。研究表明,改进的方法能很好地预测巴山冷杉种内和种间的竞争强度。     

Effects of leaf age,elevation and light conditions on photosynthesis and leaf traits in saplings of two evergreen conifers,Abies veitchii and A. mariesii

亚高山的针叶树物种分布在广泛的海拔范围内,因此它们必须具备抵抗严酷自然条件(如高海拔和长期的黑暗)的能力。 在日本中部的亚高山地带,两种常绿的针叶树冷杉(Abies veitchii)和马尾冷杉(Abies mariesii)分别在低海拔和高海拔地区占据优势。本研究的目的是探讨叶龄、海拔和光照条件对两个物种叶片形态和生理性状的影响。我们以两种针叶树的幼树为样本,分别在两种针叶树下部 (1600米)和上部(2300米)的林下层和冠层间隙中取样,并分析叶龄、海拔和光照条件对冷杉和马尾冷杉的光合作用和叶片性状的影响。研究结果表明,两种针叶树的光合作用和叶片性状对叶龄呈现相似的响应,但其对海拔和光照条件则表现出不同的响应。冷杉最大光合速率与单位叶面积质量和非结构性碳水化合物浓度呈负相关。两种树木的单位叶面积质量在高海拔处均增加,而非结构性碳水化合物浓度仅在冷杉中增加。因此,冷杉的最大光合速率在高海拔时降低。此外,两个物种的最大光合速率均与氮浓度呈正相关。在林下层,叶片氮浓度在冷杉中下降,而在马尾冷杉中上升。在林下条件下,只有马尾冷杉的单位叶面积质量下降、叶绿素-氮比增加,表明在黑暗条件下,马尾冷杉比冷杉有更高的捕光效率。本研究表明,与冷杉相比,马尾冷杉的光合和叶片特性具有更强的耐阴性,光合速率受海拔的影响较小,从而使马尾冷杉在林下植被中生存,并在高海拔地区占据优势。     

川西北甘南云冷杉林的DCA排序、环境解释和地理分布模型的研究

 本文应用DCA排序的方法进行了四川西北部和甘肃南部云冷杉林的梯度分析,建立了植物群落梯度环境解释的数学模型和植物群落地理分布的数学模型。研究结果表明：影响川西北甘南云冷杉林植物群落地理分布的生态梯度中最显著并起主导作用的是温度梯度和水分梯度。在水分梯度上,大致有这样的植被序列,由较湿的藓类和杜鹃冷杉及云杉林,中生性的箭竹冷杉林和云杉林,偏旱的高山栎冷杉林和云杉林;处于恢复中期的桦木林的生境也偏旱,但一般不如高山栎林。冷杉林比云杉林更趋于阴湿的生境。 在温度梯度上,呈现出灌木云杉林—桦木林—落阔冷杉林—高山栎云杉林—箭竹云杉林—箭竹红桦林—高山栎冷杉林—藓类红桦林—藓类云杉林—箭竹冷杉林—藓类云冷杉林—藓类冷杉林—小叶章云杉林—杜鹃冷杉林的排序。基本特征是：川西北和甘南的云杉林的温度需求高于冷杉林。川西北云冷杉林与甘南云冷杉林在DCA排序轴上没有明显的区别。虽然川西北地区云冷杉林的海拔分布高于甘南地区,但是其纬度也低于甘南,故在温度条件上基本一致。在水分梯度上,川西北云冷杉林的湿度要稍大于甘南云冷杉林。利用植物地理分布的数学模型和有关的图、表,详细地分析了植物群落的空间分布,进行了环境解释,并找出了主导的环境因子。     

峨眉山冷杉种群研究

依据 5 1个 2 0 m× 2 0 m冷杉乔木层样方和 1 0 2个 4 m× 4 m灌木层样方有关冷杉种群的调查资料和典型区域内冷杉种群的年龄和生长指标的测定 ,对峨眉山冷杉种群配置、重要参数、更新与生长状况进行了研究 ,揭示了当地冷杉种群及其所组成的森林的现状与演替趋势。指出 ,峨眉山冷杉种群的径级配置与海拔高度为主导的环境要素变化相关 ,2 80 0 m以下的冷杉种群缺乏胸径 5～ 2 0 cm的小径个体 ;尤其是在“混交”状的冷杉林下株高小于 5 m的幼龄个体稀少且其年龄与应有的生长量相差甚远 ,由此作者认为该海拔段以下的冷杉种群正面临衰退 ,其构成的冷杉森林正处于退化演替的过程之中 ,并有可能在今后 5 0 a或许更短的时间内显现严重后果。同时 ,本研究表明冷杉主要通过林窗和林缘更新。通过对冷杉 -箭竹 -泥炭藓森林群落中冷杉种群状况的研究 ,阐述了冷杉种群在特定的逆境条件下生殖策略由 K选择向 r选择方向的转化趋势 ,讨论了冷杉种群在泥炭藓发育的局部沼泽化的环境中得以生存的机制 ;本文还就冷杉种群在林内自然更新和迹地更新条件下的胸径生长规律作了比较研究 ,说明密度制约机制对冷杉种群更新具有重要作用 ;并就更新情况相对较好的冷杉 -箭竹 +金顶杜鹃 -草类林、冷杉 -箭竹 -藓类林和冷杉 -峨眉玉山     

青藏高原东缘红桦-岷江冷杉次生林中大径级保留木对其他林木的影响

为了研究森林采伐后保留木在次生群落恢复过程中对其他林木的影响, 采用Hegyi单木竞争指数(CI)和标记相关点格局k_d(r)函数对4 hm ²红桦-岷江冷杉(Betula albosinensis-Abies faxoniana)次生林样地中大径级保留木与其他林木的空间关系进行了分析。结果表明: 1)优势树种红桦(Betula albosinensis)与岷江冷杉(Abies faxoniana)的种间、种内竞争中, 种间Hegyi单木竞争指数(CI) (2.484) >红桦种内CI (1.711) >岷江冷杉种内CI (1.548); 2)大径级保留木与其他径级林木间的竞争指数中, 红桦大径级保留木与红桦小树的竞争指数最大, 红桦大径级保留木与岷江冷杉小树、岷江冷杉大径级保留木与红桦小树和岷江冷杉小树这三组的竞争指数之间无显著差异, 且三者的值最小; 3)在14.5-15.5 m尺度上, 红桦大径级保留木与岷江冷杉幼树呈显著的空间正关联; 在6.5-7.5 m的尺度上, 岷江冷杉大径级保留木与岷江冷杉幼树呈显著的空间负关联; 0-100 m尺度上, 岷江冷杉幼苗(树龄≥4年)与两种大径级保留木均呈相互独立的空间关系。     

Crossability relationships between Korean firs Abies koreana , A. nephrolepis and A. holophylla and some other representatives of the genus Abies

Artificial crossing experiments involving 3 Abies species native to Korean peninsula and 5 other representatives of Abies revealed a high hybridological affinity between Abies koreana and A. nephrolepis. Both these species are reproductively isolated from A. holophylla. All the three Korean species were found to exhibit incompatible relationships with the North American species A. concolor. The species A. holophylla and A. koreana differ also in their abilities to intercross with the Mediterranean firs. The former has been successfully crossed with A. nordmanniana, A. alba and A. cilicica exhibiting 19.1–55.3% crossability, whereas the latter produced filled seeds only with A. nordmanniana reaching 46.4% crossability. A considerable differentiation is postulated to exist between the pair of species A. koreana and A. nephrolepis on the one side and A. holophylla on the other side.     

Development and characterization of microsatellite markers in Abies firma and interspecific amplification in other Japanese Abies species

Abies firma is a dominant coniferous tree species endemic to Japan. We isolated eight microsatellite loci from needles of this tree species and tested their polymorphism among 26 A. firma individuals. Six of them showed polymorphism, with two to 16 alleles per locus. Their expected heterozygosities ranged from 0.075 to 0.922. Moreover, interspecific amplification among Abies sachalinensis, Abies mariesii and Abies veitchii was successful in majority of the isolated loci, suggesting that these loci may be useful for characterization of other fir species.     

冷杉属植物叶角质层内表面的观察及其系统学意义

在扫描电镜下,比较观察了冷杉属全世界36种2变种的叶角质层内表面的结构特征,根据叶角 质层内表面胞间凸缘特征的显著差异,将冷杉属植物归为4类:(1)直且发达的单凸缘;(2)双凸缘; (3)不发达的单凸缘;(4)弯曲且发达的单凸缘。并发现这四种凸缘类型在冷杉属中的分布格局与该 类植物的地理分布具有相关性。根据所观察的结果和前人的属下分类系统以及化石资料,认为北美西部是现代冷杉属植物的一个多样性中心。     

Molecular phylogenetic position of Japanese Abies (Pinaceae) based on chloroplast DNA sequences

The phylogenetic positions of Japanese Abies species (A. firma, A. homolepis, A. veitchii, A. sachalinensis, and A. mariesii) were revealed based on nucleotide sequences of chloroplast DNA. First, rbcL sequences of 24 Abies species worldwide were analyzed to clarify the phylogenetic position of the Japanese Abies within this genus. While 4 of the Japanese species formed a monophyletic group with almost no sequence divergence, A. mariesii was placed into a clearly different group with North American species. Second, to clarify the relationships among the species in the monophyletic group, sequences of rbcL, matK, and six spacer regions (total ca. 5 kb) were analyzed for 12 species of the group, with A. mariesii as an outgroup. Although a unique sequence and some species-specific sequences were detected in A. mariesii and in A. firma and A. homolepis, respectively, A. veitchii and A. sachalinensis gave identical sequences for all the sequenced regions.     

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.     

Morphological diversity and structure of West Mediterranean Abies species

The aim of this study was the biometrical comparison of Abies pinsapo Boiss., Abies maroccana Trab. and Abies tazaotana Côzar ex Villar from Spain and Morocco. The morphological and anatomical analysis of the needle characters of these firs revealed significant taxonomic differences. The results showed considerable distances between regions and very small differences among the Moroccan fir taxa. The examination of the needles and earlier genetic analysis did not provide evidence for distinguishing A. tazaotana at the species level.     

Microfungi associated with Abies needles and Betula leaf litter in a subalpine coniferous forest

We investigated microfungal assemblages on leaf litter within a subalpine forest in central Japan and their variation with season, litter depth, and litter species. Microfungal assemblages were compared for Abies needles and Betula leaf litter collected from litter and fermentation layers of the forest floor during the growing season in spring, summer, and autumn. A total of 35 and 42 species were isolated from Abies needles and Betula leaf litter, respectively. The observed variation in microfungal assemblages was primarily attributable to seasonal differences. The frequencies of Trichoderma viride, Volutella ciliata, Mucor sp., and Umbelopsis ramanniana increased in summer, leading to a high degree of similarity of microfungal assemblages in different litter depths and litter species. Microfungal assemblages on Abies needles in spring and autumn and those on Betula leaves in spring were characterized by Trichoderma viride, V. ciliata, Thysanophora penicillioides, Trichoderma polysporum, and (or) Mortierella alpina. Microfungal assemblages on Betula leaves in autumn were characterized by the absence of these species and the occurrence of Cladosporium cladosporioides. The results were discussed with an emphasis on the role of microfungi in decomposition processes and the impact on fungi of predicted future increases in global temperature.     

Genetic variation at chloroplast microsatellites (cpSSRs) in Abies nebrodensis (Lojac.) Mattei and three neighboring Abies species

Abies nebrodensis (Lojac.) Mattei (Sicilian fir) is an endangered species represented by only one population of 29 adult individuals occurring in a limited area of the Madonie Range in northern Sicily (Italy). Taxonomic boundaries between this taxon and the neighboring Abies species are not clear. In this study, we used six chloroplast simple-sequence repeats (cpSSRs) to investigate the population genetic structure and the distribution of chloroplast haplotypic variation in A. nebrodensis and three of the neighboring Abies species: Abies alba (Mill.), Abies numidica (De Lann) and Abies cepha-lonica (Loud.). Our aims were to quantify the level of cpDNA differentiation within the Abies populations and to shed light on the history of A. nebrodensis. Diversity levels based on the haplotype frequency at six cpSSRs were high, especially in A. alba and A. cephalonica. In all, we found 122 haplotypes among the 169 individuals analyzed, and the four species were distinguished from each other by their haplotype composition. The majority of the haplotypes (76%) were detected only once, but in A. nebrodensis seven individuals (41% of the sample population) shared the same haplotype. Moreover, the seven A. nebrodensis individuals with an identical haplotype showed a tendency to be geographically grouped within the population"s limited range. The analysis of molecular variance (AMOVA) showed a significant difference in the level of apportionment of gene diversity between the species A. alba and A. cephalonica (F_ST=0.191 and 0.012, respectively). AMOVA analysis conducted over all populations from the four species showed that 19% of the total cpSSR variation was attributable to differences among species, 6% was due to differences among populations within species, and 74% to differences within populations. The high percentage of unique haplotypes identified confirms the power of cpSSR haplotype analysis for identifying individual trees in individual Abies populations. Our results indicate that A. nebrodensis differs from the other three Abies species investigated and support its classification as an independent taxon. The results also showed a decreased level of variation in A. nebrodensis and suggested that the species has experienced a genetic bottleneck during the last two centuries. Received: 10 April 2000 / Accepted: 13 July 2000     

Chloroplast DNA inversion polymorphism in populations of Abies and Tsuga

Polymorphism for a 42-kb chloroplast DNA inversion was detected in five species of Abies and two species of Tsuga based on a sample of 1,281 individuals and both Southern hybridization and polymerase chain reaction (PCR) analyses. Two haplotypes were observed in all populations and species. The 42-kb inversion is associated with a short inverted repeat that includes trnS, psaM, and trnG. The frequencies of the two haplotypes within species were very similar among the five species of Abies This polymorphism has been maintained within populations and species in both Abies and Tsuga, probably because the mutation rate of the inversion is high. Haplotype frequencies had no geographical tendencies for any species except Abies mariesii, in which haplotype frequencies varied clinally, possibly as a result of rapid dissemination after the most recent glacial period and random genetic drift.