欧洲水青冈(Fagus sylvatical L.)构筑型与形态多样性研究

通过对植物构件和构筑型的分析,可以了解植物的整体结构与系统演化关系。高等植物构筑型的研究以树木最为深入,植物学家对全球的树木进行了构筑模式的分类,并确定了２３个基本的构筑类型。植物的构筑型和其形态的多样性是密不可分的。构筑奕型确定了植物所属的整体形态类型,而对每种植物具体的形态多样性分析可以深入了解每种构筑型的数量特征及其形成过程。在概述了树木树筑型研究的基础上,系统说明了欧洲水青冈构筑才形态多样性分析的步骤和步骤和分析的意义。水青冈属（Fagus)为Ｔroll型构筑模式,处于不同生态条件下的同一种的不同种群之间,在形态、解剖、生理和遗传多样性方面都存在着差异。树木的形态多样性虽然主要受遗传因素的控制,但生态条件对其形态和遗传多样性也有非常显著的影响。在对树木进行构筑型分析时,道德要根据构筑要素（主要是形态与生长特性）确定它所属的构筑类型,其次是分析它的形态特征与环境条件的关系,进行进一步分析其遗传多样性。     

The evolutionary history of Fagus in western Eurasia: Evidence from genes, morphology and the fossil record

 Fagus (beech) is among the most abundant and economically important genera of broad-leaved trees in northern hemisphere temperate forests. The number of modern taxa present in Europe and Asia Minor has long been a matter of debate and up to five species have been recognised. To resolve taxonomic and phylogenetic relationships we conducted morphological and molecular genetic analyses in western Eurasiatic taxa and evaluated palaeontological evidence. To place our findings from western Eurasiatic populations in a broader context additional East Asiatic and North American species of the same subgenus Fagus as well as two species of the subgenus Engleriana were included in our study. The morphological features exhibited in western Eurasiatic populations of Fagus show a west-east gradient that is characterised by strongly overlapping variability between geographical races. Fagus populations from Asia Minor exhibit an even higher variability that is also reflected in their genetic variability of nuclear rDNA internal transcribed spacer (ITS) sequences. The intraspecific genetic variability recorded here is in conflict with previous ITS studies in Fagus. The high amount of ITS polymorphism within Fagus from western Eurasia along with the clinal variation observed for morphological characters suggest the presence of only a single species, Fagus sylvatica L., in Europe and Asia Minor. Previously recognised taxa such as F. orientalis Lipsky and Fagus moesiaca (Maly) Czeczott should therefore be treated as synonyms of Fagus sylvatica. Although species belonging to the subgenus Engleriana were genetically distinct from species of the subgenus Fagus, relationships within the subgenus Fagus could not be clearly resolved. A reason for this could be the low rate of diversification in Fagus during the early phase of range expansion of the genus in the Oligocene period as indicated by the uniformity of leaf and cupule/nut fossils. This may account for the low overall ITS divergence and the high degree of polymorphism encountered in the subgenus Fagus and points to a late differentiation of western Eurasiatic and eastern Asiatic species. Area disruptions during the Pleistocene and the post-glacial recolonisation of western Europe appear to have caused the west-east gradient that is apparent in modern Fagus of western Eurasia but absent in Late Tertiary ancestors of Fagus sylvatica. Received June 22, 2001 Accepted February 25, 2002     

Interpretation of seasonal changes of xylem embolism and plant hydraulic resistance in Fagus sylvatica

The annual course of xylem embolism in twigs of adult beech trees was monitored, and compared to concurrent changes of tree water status and hydraulic resistances. Xylem embolism was quantified in 1-year-old apical twigs by the hydraulic conductivity as a percentage of the maximum measured after removal of air emboli. Tree and root hydraulic resistances were estimated from water potential differences and sap flux measurements. The considerable degree of twig embolism found in winter (up to 90% loss of hydraulic conductivity) may be attributed to the effect of freeze-thaw cycles in the xylem. A partial recovery from winter embolism occurred in spring, probably because of the production of new functional xylem. Xylem embolism fluctuated around 50% throughout the summer, without significant changes. Almost complete refilling of apical twigs was observed early in autumn. A significant negative correlation was found between xylem embolism and precipitation; thus, an active role of rainfall in embolism reversion is hypothesized. Tree and root hydraulic resistances were found to change throughout the growing period. A marked decrease of hydraulic resistance preceded the refilling of apical twigs in the autumn. Most of the decrease in total tree resistance was estimated to be located in the root compartment.     

Mechanism of freeze-induced embolism in Fagus sylvatica L.

 The mechanism of freeze stress-induced embolism in Fagus sylvatica L. branches was analyzed under controlled conditions. Excised branches were exposed to successive freeze-thaw cycles in temperature controlled chambers. Thermocouples were placed on the bark to detect sap freezing exotherms. The degree of xylem embolism was estimated after each cycle by the loss of hydraulic conductivity. After one freeze-thaw cycle the degree of embolism was found to decrease with xylem specific hydraulic conductivity, small apical shoots being more susceptible to embolism. Exotherms revealed that distal shoots were freezing first and exuded sap as a result of water expansion. The lower water content in apical shoots upon freezing probably induced higher sap tensions which promoted air bubble expansion and vessel cavitation preferentially near the apices. When the decrease in water content was experimentally prevented, embolism developed to a lesser extent. The higher vulnerability of shoot apices may protect the rest of the branch from winter damage. Received: 29 May 1998 / Accepted: 15 August 1998     

Genetic variation of European beech (Fagus sylvatica L.) along an altitudinal transect at Mount Vogelsberg in Hesse, Germany

Allelic and genotypic variation at 13 different enzyme loci of autochthonous European beech (Fagus sylvatica L.) was investigated in six 110-160-year-old stands growing at elevations between 150 and 660 m above sea level on the western slope of mount Vogelsberg in central Germany. The highest elevated population showed the highest number of effective alleles (Ne), the highest total heterozygosity (He) and the highest population differentiation deltaT. Also, the genotype SKD-A2A3 of shikimate dehydrogenase was significantly more frequent at the two highest elevated stands (P = 11%) than at the three lowest elevated stands (P = 1%). Further differences in genotype frequencies between 11 of 15 stand pairs were elevation independent.     

4种形态类型诸葛菜的等位酶变异及遗传多样性分析

以采自江苏省南京市4个样点的4种不同形态类型(紫花毛果、紫花光果、白花光果和白花毛果)诸葛菜〔Orychophragmus violaceus(L.)O.E.Schulz〕63个单株为研究对象,采用不连续聚丙烯酰胺凝胶电泳技术(SDS-PAGE)对过氧化物酶同工酶(POD)、乙醇脱氢酶(ADH)、谷氨酸脱氢酶(GDH)、超氧化物歧化酶(SOD)和多酚氧化酶(PPO)酶谱进行了分析,并基于5个等位酶的酶谱分析结果对4种形态类型诸葛菜的遗传多样性进行了研究。结果表明:5个等位酶系统共包含12个位点32个等位基因,其中POD-1、POD-4、POD-5、POD-6、ADH-1、PPO-1、SOD-1和SOD-2为多态性位点;除具有共有谱带外,不同类型诸葛菜还具有各自特有的酶谱特征,其中,GDH-2的b带和c带分别是紫花类型和白花类型的特有谱带。紫花毛果、紫花光果和白花毛果类型的多态位点百分率(PPL)均为58.33%,而白花光果的PPL为66.67%;紫花毛果、紫花光果、白花光果和白花毛果每个位点的平均等位基因数分别为2.42、2.25、2.42和1.83,平均期望杂合度分别为0.35、0.30、0.38和0.29。4种形态类型诸葛菜的总基因多样度平均值为0.61,类型内基因多样度平均值为0.49,明显大于类型间基因多样度平均值(0.12);类型间的基因分化系数平均值为0.195,表明总基因多样性的80.5%来源于类型内。紫花光果和白花毛果类型间的遗传一致度最低(0.724 4)且遗传距离最大(0.322 5),而2个白花类型间的遗传一致度最高(0.954 1)且遗传距离最小(0.047 1)。研究结果显示:诸葛菜的种内变异程度很大、遗传分化程度较高,但在各类型内具有较高的遗传相似性。     

Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica)

Aim  Establishing possible relationships between the magnitudes of the glacial distribution of the European beech, Fagus sylvatica L., and its post-glacial spread.
Location  Europe.
Methods  A database of over 400 pollen records has been used to locate Fagus populations at the end of the last glacial and during the post-glacial in Europe and to assess the areal extent of their past distribution.
Results  The rate of late-glacial and post-glacial increase in the number of pollen sites where Fagus was locally present conforms well to a logistic model of population growth. This suggests that the area occupied by beech populations expanded exponentially from the glacial refugia for a duration of over 10,000 years, until about 3500 yr bp . In the past three millennia beech populations increased at a slower rate, tending towards an equilibrium value.
Main conclusions  The conformity of the increase in beech distribution to the classical logistic model of population growth indicates that: (1) a multiplicative biological process was the main factor shaping the pattern of the post-glacial expansion of F. sylvatica in Europe, (2) climate conditions, human activity and competition may have influenced its rate of spread, and (3) beech populations did not expand with a moving closed front, but with a diffuse spread from scattered nuclei. The distribution of Fagus in Europe at the end of the last glacial appears to have been of two orders of magnitude less extensive than at present. Pleistocene refugia were likely to have been a mosaic of sparse stands of small populations scattered in multiple regions. Fagus populations appear to have increased very slowly and to a moderate extent in southern Europe, where they are now declining slightly. The central European populations increased quickly and extensively, reaching northern Europe, and are now approaching their carrying capacity.     

Development of protein bodies in cotyledons of Fagus sylvatica

Electron microscopic analysis of cotyledons of beech ( Fagus sylvatica ) at different stages of seed maturation indicates that protein bodies originate by gradual subdivision of the vacuoles in which reserve proteins are deposited. The majority of protein bodies show a proteinaceous matrix and a number of globoid inclusions of different sizes. In a small number of cells, druse-like inclusions were observed. Analysis by SDS-PAGE of proteins extracted from isolated protein bodies shows that the majority correspond to beech seed storage globulins.     

Analysis of natural Fagus sylvatica L. s.l. regeneration in low elevation stands located in the central part of the Evros region in northeastern Greece: Is sprout origin regeneration significant for species maintenance?

Abstract

The aim of this study was (a) to analyse natural Fagus sylvatica L. s.l. regeneration in low elevation stands located in the central part of Evros region in northeastern Greece in relation to factors such as site productivity, type of silvicultural treatment and regeneration origin and (b) to determine whether or not sprout origin regeneration is significant for the maintenance of beech stands. One hundred and twenty plots (3 m × 3 m) were established in areas where thinnings and regeneration fellings had taken place, and in two site productivity regimes. In each plot, the number of beech seedlings, saplings, and stump or root sprouts (regeneration plants) with a height of up to 3 m was counted, and their height measured. Our findings indicate that even though reproduction by seed is the predominant regeneration mechanism in our stands, in the medium productivity sites sprouts represent a significant percentage of the total number of plants. Moreover, sprouting functions as a mechanism of maintenance of beech stands in medium productivity sites. During the regeneration fellings practised in this area, it is advisable to cut the seedlings and saplings of the advanced regeneration in order to supplement the seed-derived plants with seedling sprouts.     

Characterization of microsatellite markers in Fagus sylvatica L. and Fagus orientalis Lipsky

Using an enrichment procedure, we cloned microsatellite repeats from European beech (Fagus sylvatica L.) and developed primers for the amplification of microsatellite markers. Six polymorphic loci were characterized which produced 3–21 alleles in 70 individuals from one Italian population, with an observed heterozygosity between 0.58 and 0.85. All six loci amplified fragments which were polymorphic in the closely related species, Fagus orientalis, also. Owing to their very high degree of variation, these markers should be very useful in gene flow studies of these species.     

The effect of shoot architecture on hydraulic conductance in beech (Fagus sylvatica L.)

 Shoot hydraulic conductance was measured in beech (Fagus sylvatica L.) that had previously been exposed to high levels of nitrogen input. Whole-shoot hydraulic conductance, conductance per unit pressure gradient and leaf specific conductance were negatively correlated with the number of bud scars per unit length, a morphological parameter of tree decline. We propose a negative feedback mechanism by which stress induced alterations in shoot morphology can cause a lasting reduction of tree vigour. Received: 2 July 1997 / Accepted: 25 June 1998     

The variability of vessel size in beech (Fagus sylvatica L.) and its ecophysiological interpretation

The vessel areas of ten beech trees growing on a dry site were measured separately for all tree rings using automatic image analysis. These data were correlated with the monthly amount of precipitation from the July prior to the growing season until the August of the current growing season. It is evident that vessel formation at the beginning of cambial activity is mainly controlled by internal factors. The rainfall in the previous summer and autumn and in the contemporary May had only a slight influence. Vessel formation towards the end of the cambial activity is strongly influenced by the July rainfall and is thus determined to a greater degree by external factors. These results are discussed on the basis of hypotheses of tree physiology.     

Spatial Distribution of Genetic Variation in a Natural Beech Stand (Fagus sylvaticaL.) Based on Microsatellite Markers

The spatial distribution of alleles is described in a naturally regenerated, isolated pure beech (Fagus sylvaticaL.) stand consisting of 99 adult trees. After testing nine microsatellite loci originally developed for F. crenata, each tree was genotyped at four well-scorable microsatellite loci. Specific primers were developed for one locus of F. sylvaticaL. For the characterization of spatial genetic structures, two different statistics were used. One method is based on the mean genetic distance between trees in different spatial distance classes, and the other one is Moran's index I. The results show the same tendency of a strong family structure in the distance classes up to 30m in comparison with that expected for a spatially non-systematic distribution of genotypes. In general, microsatellites are more useful to detect spatial genetic structures than allozymes. Spatial genetic structures are influenced by unpredictable factors such as wind direction at anthesis and can therefore vary from year to year. We recommend that seed collections should cover large areas in order to prevent a preponderance of few families and a reduction of the adaptive potential of the next generation.     

Storage and remobilisation of sulphur in beech trees (Fagus sylvatica)

Three-year-old beech trees were fed ³⁵S-sulphate in August 1993 via a flap in a mature leaf of an upper branch. Harvest of beech trees was performed 24 h after feeding ³⁵S-sulphate, before leaf senescence, after leaf abscission, in early winter (January 1994). in late winter (March 1994). before bud break and after bud break. Twenty-four h after feeding ³⁵S-sulphate, 0.7 ± 0.5% of the ³⁵S-radioactivity taken up was exported out of the fed leaf. When trees were analysed 2 months later, i.e., before leaf senescence, this value had increased to 22 ± 7%. The exported ³⁵S-radioactivity was located in the branch containing the fed leaf (2.8 ± 13%). in basipetal parts of the trunk (41 ± 77%) and in the main rool (21 ± 6%). Leaves and apical parts of the trunk were no sink organs for the exported sulphur. Along the tree axis the main proportion of the radiolabel was located in the wood, predominantly in the acid soluble fraction. In the bark the greater portion of the radiolabel was found in the acid insoluble fraction. In both tissues the bulk of the ³⁵S of the soluble fraction was sulphate together with small amounts of glutathione. This pattern did not change until bud break. After bud break, basipetal parts of the trunk lost part of its ³⁵S-radioactivity. Of the ³⁵S-radioactivity which had been exported out of the fed leaf during the previous autumn, 16 ± 2% remained in the trunk, whereas 47 ± 7% of the ³⁵S was found in branches, mainly in the newly developed leaves. The present results show that sulphur, mainly in the form of sulphate, is stored along the tree axis in both bark and wood of beech trees and is re-mobilised during leaf development in spring.     

Assessment of genetic relationships and population discrimination among Fagus sylvatica L. by RAPD

 We assessed the genetic relationships between members of the Fagaceae family by RAPDs in order to better ascertain the taxonomic status of a very particular population of Fagus sylvatica, the ‘tortuosa’ variety. Intra- and inter-population Nei and Li’s mean genetic distances were compared, and the genetic relationships between individuals were clarified on dendrograms by the Neighbor joining method. RAPD analysis was first conducted on three species from three genera, Quercus petraea, Castanea sativa, and Fagus sylvatica, in order to develop an efficient RAPD protocol. The variety level was then studied, and a general tendency of the individuals to cluster by variety was observed. Individuals also clustered by geographic locations, but the genetic distances between populations were not correlated to the distances between sites. Finally, we compared the common beech and ‘tortuosa’ varieties from two different locations, Verzy and Süntel. Both populations from one location were closer than the same variety from two sites. This last result is in agreement with those previously obtained with isozymes. Hypotheses concerning the origin of the ‘tortuosa’ variety are discussed. Received: 9 January 1998 / Accepted: 25 February 1998     

Patterns of genetic variation in Pacific island land snails: the distribution of cytochrome b lineages among Society Island Partula

The radiation of Partula land snails has produced a large array of distinct morphological, ecological and behavioural types occupying many tropical volcanic islands in the Pacific Ocean. Within the Society Islands of French Polynesia, the mode of evolution is thought to have involved a single colonization event on each island, with later speciation occurring largely in situ. The present study examines genetic variation in the mitochondrial cytochrome b gene among taxa within the Society Island archipelago. Levels of intraspecific variation are found to be high, but variation among species is sometimes small. Mitochondrial variants do not always cluster according to island and some species are found to be polyphyletic in the cytochrome b tree, despite other morphological and molecular evidence that strongly supports their monophyly. A possible explanation for the polyphyly of species is that different variants are derived from ancestral mitochondrial polymorphisms that have been retained despite speciation events. Although it is possible that there has been some gene flow among islands, the distribution of mitochondrial lineages across islands strongly indicates that their origins predate the colonization of the islands in the study, and that they are very unlikely to have evolved entirely in situ.     

Local differentiation of Albanian provenances and chorology of beech species in balcan area

Abstract

Some quantitative characters of different Albanian beech provenances are analysed in order to study their local differentiation. The results obtained by discriminant analysis showed the formation of two principal groups. Biza provenance was discriminated from both groups. Principal components and cluster analysis give a good evidence on the similarities among Albanian and Bulgarian beech populations. The results suggest that all area of Balcan is the contact zone of Fagus sylvatica L. and Fagus orientalis Lypski.