Hybridization and polyploidy as drivers of continuing evolution and speciation in Sorbus

Interspecific hybridization and polyploidy are pivotal processes in plant evolution and speciation. The fate of new hybrid and polyploid taxa is determined by their ability to reproduce either sexually or asexually. Hybrids and allopolyploids with odd chromosome numbers are frequently sterile but some establish themselves through asexual reproduction (vegetative or apomixis). This allows novel genotypes to become established by isolating them from gene flow and leads to complex patterns of variation. The genus Sorbus is a good example of taxonomic complexity arising from the combined effects of hybridization, polyploidy and apomixis. The Avon Gorge in South‐west Britain contains the greatest diversity of Sorbus in Europe, with three endemic species and four putative endemic novel hybrids among its 15 native Sorbus taxa. We used a combination of nuclear microsatellite and chloroplast DNA markers to investigate the evolutionary relationships among these Sorbus taxa within the Avon Gorge. We confirm the genetic identity of putative novel taxa and show that hybridization involving sexual diploid species, primarily S. aria and S. torminalis and polyploid facultative apomictic species from subgenus Aria, has been responsible for generating this biodiversity. Importantly our data show that this creative evolutionary process is ongoing within the Avon Gorge. Conservation strategies for the rare endemic Sorbus taxa should therefore consider all Sorbus taxa within the Gorge and must strive to preserve this evolutionary process rather than simply the individual rare taxa that it produces.     

花楸属3种植物的基因组大小与叶气孔特征分析

以糍粑沟花楸(Sorbus cibagouensis H.Peng&Z.J.Yin)、大理花楸(S.hypoglauca(Cardot)Hand.-Mazz.)和川滇花楸(S.vilmorinii C.K.Schneid.)为材料,采用流式细胞术对其基因组大小及倍性进行检测分析,同时应用光学显微镜和扫描电子显微镜对其气孔特征进行观察。结果显示,3种花楸属植物的基因组大小和倍性、气孔特征均存在一定差异。糍粑沟花楸、大理花楸和川滇花楸的基因组大小分别为:(1.480±0.039)pg、(1.513±0.041)pg、(2.675±0.065)pg,在此基础上推断糍粑沟花楸和大理花楸为二倍体、川滇花楸为四倍体植物。显微镜观测发现:3种花楸属植物的气孔器均分布于叶的下表皮,气孔不下陷,保卫细胞无“T”型加厚结构,气孔类型为无规则形;糍粑沟花楸和川滇花楸的气孔器外拱盖光滑,而大理花楸气孔器外拱盖具有短棒状蜡质纹饰;3种植物的气孔器大小存在极显著差异。研究结果表明花楸属植物的基因组大小与倍性呈显著正相关,可用于推断植物的倍性;而气孔器大小和密度与倍性的相关性不大,但气孔特性在种间变化显著,可为种的鉴定提供科学的理论依据。     

花楸树种源间表型性状的地理变异分析

为揭示花楸树[ Sorbus pohuashanensis (Hance) Hedl.]表型性状的地理变异规律,对来源于山东、山西和河北的7个花楸树种源171个家系2年生幼苗的株高、地径、复叶长、复叶宽、叶柄长、小叶长、小叶宽、复叶长宽比和小叶长宽比9个表型性状进行了比较,并进行了方差分析、主成分分析(PCA)及聚类分析;在此基础上,分析了各种源株高和地径与主要地理-气候因子的相关性.结果表明:在种源间及种源内家系间 9个表型性状均有极显著差异;7个种源幼苗的平均株高和平均地径变幅分别为54.21～ 63.07 cm和8.12 ～8.45 mm;叶片各项指标的变幅相对较小,变幅最大的为叶柄长,为1.87 ～3.05 cm;9个性状在种源间的表型分化系数为12.98％～54.04％,其中小叶长的表型分化系数最大(54.04％),株高的表型分化系数最小(12.98％),种源间各性状的平均表型分化系数为34.38％.主成分分析结果表明:花楸树表型性状之间的差异主要表现在小叶长和宽,其次是复叶长和宽.通过聚类分析可将7个种源划分为2组,其中,山东崂山和泰山种源聚为I组;山西庞泉沟及河北白石山、驼梁山、雾灵山和塞罕坝5个种源聚为Ⅱ组.相关性分析结果表明:花楸树株高与纬度呈显著负相关、与降雨量呈显著正相关(P=0.05);地径则与经度、年均温和年降雨量呈显著正相关(P=0.05).研究结果显示:花楸树的各项表型性状在种源间以及种源内家系间存在丰富的变异、遗传分化显著,其中种源内家系间的变异是花楸树表型变异的主要来源.初步判断花楸树表型性状的变异为经纬向双重变异模式,其中纬度起主要作用.     

干旱胁迫下石灰花楸幼苗叶片的解剖结构和光合生理响应

采用盆栽控水设置4种水分梯度,研究1年生石灰花楸幼苗叶片解剖结构和光合生理指标对干旱胁迫的响应。结果显示:(1)干旱胁迫下,石灰花楸叶片逐渐变薄,轻度和中度胁迫下栅海比显著升高,而重度胁迫下显著降低。(2)干旱胁迫抑制了光合色素合成,降低了叶绿素和类胡萝卜素含量,同时使叶绿素a/b和叶绿素/类胡萝卜素值升高。(3)随干旱程度加剧,净光合速率、蒸腾速率和气孔导度日变化整体下降,胞间CO2浓度整体上升,光合限制以非气孔因素为主。研究表明,石灰花楸能根据水分亏缺程度调整叶片结构和光合生理特征以维持生存和生长,具有较强的耐旱性。     

花楸树种子散布、萌发与种群天然更新的关系

花楸树是我国东北林区重要的非木质资源树种,其实生天然更新不良.本文通过研究花楸树种子散布、土壤种子库及种子萌发出土过程,分析花楸树实生天然更新的影响因素.结果表明：自然散落的花楸树果实96.1%分布于母株2 m范围内,凋落物层和土壤表层（0～2 cm）的种子数占土壤种子库总数的97.0%;不同季节花楸树土壤种子库种子数量差别很大,当年11月上旬种子数量最多,达(257.7±69.2)粒·m^-2;翌年7月下旬种子数量最少,仅为(2.9±2.9)粒·m^-2;温度不是花楸树种子萌发出土过程的限制因子,0 ℃～5 ℃时幼苗出苗率达(67.5±6.6)%,但对其出苗速率影响显著.土壤含水量为50%时,花楸树出苗率最高,达(74.7±4.2)%;含水量为60%时,幼苗死亡率最低,为(32.6±0.6)%.花楸树种子的散布格局和土壤种子库的时空分布格局影响种子的萌发出土过程,进而影响其种群的天然更新.     

黄山花楸种群遗传多样性研究

利用RAPD技术,对黄山花楸(Sorbus amabilis)自然分布区的13个自然种群的遗传多样性进行了研究,从40个10碱基随机引物中筛选出能产生稳定多态性标记的引物14个,共扩增出105个位点,其中多态性位点30个,占28．6％,应用UPGMA法和Neighbor-Joining法对遗传距离进行聚类分析构建树系图。结果表明,黄山花楸自然种群具有较低的遗传多样性,对环境变化的适应能力较差;其种群间的遗传差异与其地理分布有关;黄山花楸自身的特殊进化历史和人为砍伐以及自然灾害(火灾、病虫害等)和小种群的遗传漂变作用是黄山花楸遗传多样性水平低的主要原因,也是其濒危的主要原因。     

多蕊石灰树属应与花楸属合并

多蕊石灰树属与花楸属在性状上没有明显区别,故应予以合并。提出了墨脱花楸Sorbus medogensis 这一新名称。     

Competitive success of southern populations of Betula pendula and Sorbus aucuparia under simulated southern climate experiment in the subarctic

Global warming has been commonly accepted to facilitate species’ range shifts across latitudes. Cross‐latitudinal transplantations support this; many tree species can well adapt to new geographical areas. However, these studies fail to capture species’ adaptations to new light environment because the experiments were not designed to explicitly separate species’ responses to light and temperature. Here we tested reaction norms of tree seedlings in reciprocal transplantations 1,000 km apart from each other at two latitudes (60°N and 69°N). In contrast to past studies, we exposed our experimental plants to same temperature in both sites (temperature of 60°N growing site is recorded to adjust temperature of 69°N site in real time via Internet connection) while light environment (photoperiod, light quality) remained ambient. Shoot elongation and autumn coloration were studied in seedlings of two deciduous trees (Betula pendula and Sorbus aucuparia), which were expected to respond differently to day length. Sorbus as a member of Rosaceae family was assumed to be indifferent to photoperiod, while Betula responds strongly to day length. We hypothesized that (1) southern and northern populations of both species perform differently; (2) southern populations perform better in both sites; (3) autumn phenology of southern populations may delay in the northern site; (4) and Sorbus aucuparia is less dependent on light environment. According to the hypotheses, shoot elongation of northern population was inherently low in both species. An evolutionary consequence of this may be a competitive success of southern populations under warming climate. Southern population of B. pendula was delayed in autumn coloration, but not in growth cessation. Sorbus aucuparia was less responsive to light environment. The results suggest that light provides selection pressure in range shifts, but the response is species dependent.     

Fruit softening: evidence for pectate lyase action in vivo in date (Phoenix dactylifera) and rosaceous fruit cell walls

Background and AimsThe programmed softening occurring during fruit development requires scission of cell wall polysaccharides, especially pectin. Proposed mechanisms include the action of wall enzymes or hydroxyl radicals. Enzyme activities found in fruit extracts include pectate lyase (PL) and endo-polygalacturonase (EPG), which, in vitro, cleave de-esterified homogalacturonan in mid-chain by β-elimination and hydrolysis, respectively. However, the important biological question of whether PL exhibits action in vivo had not been tested.MethodsWe developed a method for specifically and sensitively detecting in-vivo PL products, based on Driselase digestion of cell wall polysaccharides and detection of the characteristic unsaturated product of PL action.Key ResultsIn model in-vitro experiments, pectic homogalacturonan that had been partially cleaved by commercial PL was digested to completion with Driselase, releasing an unsaturated disaccharide (‘ΔUA–GalA’), taken as diagnostic of PL action. ΔUA–GalA was separated from saturated oligogalacturonides (EPG products) by electrophoresis, then subjected to thin-layer chromatography (TLC), resolving ΔUA–GalA from higher homologues. The ΔUA–GalA was confirmed as 4-deoxy-β-l-threo-hex-4-enopyranuronosyl-(1→4)-d-galacturonic acid by NMR spectroscopy. Driselase digestion of cell walls from ripe fruits of date (Phoenix dactylifera), pear (Pyrus communis), rowan (Sorbus aucuparia) and apple (Malus pumila) yielded ΔUA–GalA, demonstrating that PL had been acting in vivo in these fruits prior to harvest. Date-derived ΔUA–GalA was verified by negative-mode mass spectrometry, including collision-induced dissociation (CID) fragmentation. The ΔUA–GalA:GalA ratio from ripe dates was roughly 1:20 (mol mol^–1), indicating that approx. 5 % of the bonds in endogenous homogalacturonan had been cleaved by in-vivo PL action.ConclusionsThe results provide the first demonstration that PL, previously known from studies of fruit gene expression, proteomic studies and in-vitro enzyme activity, exhibits enzyme action in the walls of soft fruits and may thus be proposed to contribute to fruit softening.     

Comparing direct vs. indirect estimates of gene flow within a population of a scattered tree species

The comparison between historical estimates of gene flow, using variance in allelic frequencies, and contemporary estimates of gene flow, using parentage assignment, is expected to provide insights into ecological and evolutionary processes at work within and among populations. Genetic variation at six microsatellite loci was used to quantify genetic structure in the insect-pollinated, animal-dispersed, low-density tree Sorbus torminalis L. Crantz, and to derive historical estimates of gene flow. The neighbourhood size and root-mean-squared dispersal distance inferred from seedling genotypes ( N _b  = 70 individuals, σ _e  = 417 m) were similar to those inferred from adult genotypes ( N _b  = 114 individuals, σ _e  = 472 m). We also used parentage analyses and a neighbourhood model approach after an evaluation of the statistical properties of this method on simulated data. From our data, we estimated even contributions of seed- and pollen-mediated dispersal to the genetic composition of established seedlings, with both fat-tailed pollen and seed dispersal kernels, and slightly higher mean distance of pollen dispersal (248 m) as compared to seed dispersal (135 m). The resulting contemporary estimate of gene dispersal distance (σ _c  = 211 m) was ∼twofold smaller than the historical estimates. Besides different assumptions and statistical nuances of both approaches, this discrepancy is likely to reflect a recent restriction in the scale of gene flow which requires manager's attention in a context of increasing forest fragmentation.