三种豆科灌木对干旱胁迫的响应与适应

干旱胁迫是全球范围内影响植物生存、生长和分布的重要环境因子。岷江上游干旱河谷区，由于生态环境的脆弱性和长期人类活动的干扰和过度利用，导致植被严重退化，水土流失加剧，山地灾害频繁，干旱化和荒漠化趋势明显。这种趋势若不能遏制，将严重阻碍区域社会经济的快速协调发展，并且威胁成都平原地区的发展和长江中下游地区的生态安全。因而开展干旱河谷生态恢复研究成为解决这些问题的关键。水分匮乏是限制干旱河谷生态恢复的关键因子，在全球气候变化的背景下，干旱胁迫在区域尺度上可能会更加严重，并使干旱河谷的生态环境更加恶化。因此，深入研究干旱河谷乡土植物对干旱胁迫的响应和适应机理，具有非常重要的理论和实践意义。 本论文以岷江上游干旱河谷的三种乡土豆科灌木，白刺花（Sophora davidii）、小马鞍羊蹄甲（Bauhinia faberi var. microphylla）和小雀花（Campylotropics polyantha）理论和实践意义。为研究对象，在人工控制条件下设计了4-5个连续性干旱胁迫处理，系统地研究了灌木幼苗的生长、生物量积累和水分利用效率（WUE）、形态结构和生理过程等对干旱胁迫的反应，揭示了幼苗的干旱适应能力及种间差异。主要研究结论如下： 1) 灌木生长和繁殖对干旱胁迫的反应 在干旱胁迫下，幼苗生长速率显著减小，叶片衰老和脱落比率增大，这些变化随着胁迫强度的增加具有累积效应。叶片比茎对干旱胁迫的反应更敏感。在严重干旱胁迫下，幼苗的有性繁殖被限制，但在中等程度干旱胁迫下，幼苗的有性繁殖能力被提高。 2) 灌木生物量积累及其分配和WUE对干旱胁迫的反应 在干旱胁迫下，灌木各器官的生物量都显著减小，但是生物量的分配侧重于地下部分，使得根茎比在干旱条件下增大。幼苗的耗水量（WU）随着干旱胁迫的增加而显著减少。白刺花和小马鞍羊蹄甲WUE在干旱胁迫下降低；小雀花的WUE在中等干旱胁迫下升高。 3) 灌木叶片结构特征对干旱胁迫反应 白刺花叶片具有较为典型的旱生型结构，而小马鞍羊蹄甲和小雀花则为中生型结构。在1至2年的干旱胁迫下，灌木叶片结构组成未发生本质性的改变，主要是细胞大小的变化。在中等和严重干旱胁迫下，叶肉组织厚度明显减小；并且气孔和表皮细胞面积也显著减小，气孔和表皮细胞密度显著增大；叶肉细胞层数、P/S值、表皮厚度等无显著变化。 4) 灌木对干旱胁迫的生理响应 气体交换参数和叶片相对含水量（RWC）在中等干旱胁迫下发生了明显的改变，而叶绿素荧光参数和光合色素含量在严重干旱胁迫下才发生显著变化。在干旱胁迫下，净光合作用速率（Pn）、气孔导度（gs）和RWC呈下降趋势，而叶片温度（Tl）呈增加趋势，蒸腾速率（Tr）的变化不明显。除了日最大Pn减小以外，干旱胁迫对气体交换参数的日变化无显著影响，但是对光合-光响应曲线有显著的影响，使有效光合时间缩短。在严重干旱胁迫下光系统受到损害而代谢减弱，PSⅡ中心的内禀光能转换效率（Fv/Fm）、量子产量（Yield）、光化学淬灭参数（qP）显著降低，而非光化学淬灭参数（NPQ）明显增加。气孔限制和非气孔限制对Pn的影响与干旱胁迫强度有关。在中度胁迫下，气孔限制起主导作用，在严重胁迫下非气孔限制起主导作用，40% FC水分条件可能是灌木由气孔限制向非气孔限制的转折点。 5) 灌木对干旱胁迫的适应能力及其种间差异 三种灌木对干旱胁迫具有较好的适应能力，即使在20% FC，幼苗未因干旱胁迫III而死亡；80% FC适宜于幼苗生长。白刺花生长速率慢，耗水量较少，具有较强的耐旱和耐贫瘠能力，并具有干旱忍受机制，能够在较干旱的环境中定居和生长。小马鞍羊蹄甲和小雀花，生长快，水分消耗量较大，尤其是小雀花，对干旱胁迫的忍受能力较弱，具有干旱回避机制，因而适宜于在较为湿润的生境中生长。综合分析表明，生长速率较慢的物种抗旱能力较强，其更适宜于作为干旱地区植被恢复物种。 Drought is often a key factor limiting plant establishment, growth and distribution inmany regions of the world. The harsh environmental conditions and long-termanthropogenic disturbance had resulted in habitat destruction in the dry valley ofMinjiang river, southwest China. Recently, it tended to be more severe on the vegetationdegradation, soil erosion and water loss, natural disaster, as well as desertification, whichimpact on regional booming economy and harmonious development, and would be verydangerous to the environmental security in the middle and lower reaches of Yangzi River.Therefore, ecological restoration in the dry valley is one of the vital tasks in China. Waterdeficit is known to affect adversely vegetation restoration in this place. Moreover, in thecontext of climate change, an increased frequency of drought stress might occur at aregional scale in the dry valleys of Minjiang River. The selection of appropriate plantingspecies for vegetation restoration in regard to regional conditions is an important issue atpresent and in further. The research on responses of indigenous species to drought stresscould provide insights into the improvement of the vegetation restoration in the dry valleys of Minjiang River. In this paper, the responses of three indigenous leguminous shrubs, Sophora davidii,Bauhinia faberi var. microphylla and Campylotropics polyantha, to various soil watersupplies were studied in order to assess drought tolerance of seedlings, and to compare interspecific differences in seedlings’ responses to drought stress. The results were as follows: 1 Growth and reproduction of shrubs in response to drought stress Seedling growth reduced significantly while leaf senescence accelerated underdrought stress, the cumulative responses to prolonged drought were found. The capacityfor reproduction was limited by severe drought stress, and improved by moderate droughtstress. Leaf responses were more sensitive than shoot to various water supplies. 2 WUE, biomass production and its partitioning of shrubs in response to drought stress Drought stress reduced significantly the total dry mass and their components ofseedlings, and altered more biomass allocation to root system, showing higher R/S ratiounder drought. Water use (WU) and water-use efficiency (WUE) of both S. davidii and B.faberi var. microphylla declined strongly with drought stress. The WU C. polyantha ofalso declined with drought stress, but WUE improved under moderate drought stress. 3 Anatomical characteristics and ultrastructures of leaves in response to drought stress There were xeromorphic for S. davidii leaves and mesomorphic for B. faberi var.microphylla and C. polyantha at the all water supplies. The foundational changes in leafstructures were not found with drought stress. However, mesophyll thickness, the areas ofstomatal and epidermis reduced slightly while the densities of stomatal and epidermisincreased under severe drought stress. Variations in these parameters could mainly be duoto cell size. Other structures did not displayed significant changes with drought stress. 4 Physiological responses of shrubs to drought stress The gas exchange parameters and leaf relative water content (RWC) were affectedby moderate stress, while chlorophyll fluorescence and chlorophyll content were onlyaffected by severe stress. Drought stress decreased net photosynthesis rate (Pn), stomatalconductance, light-use efficiency and RWC, and increased leaf temperature. Therespiration rates (Tr) were kept within a narrower range than Pn, resulting in aprogressively increased instantaneous water use effiecency (WUEi) under drought stress.Moreover, drought stress also affected the response curve of Pn to RAR, there was adepression light saturation point (Lsat) and maximum Pn (Pnmax) for moderate andsevere stressed seedling. However, diurnal changes of gas exchange parameters did notdiffer among water supplies although maximum daily Pn declined under severe stress.VISevere stress reduced Fv/Fm, Yield and qP while increased NPQ and chlorophyll content.Photosynthetic activity decreased during drought stress period due to stomatal andnon-stomatal limitations. The relative contribution of these limitations was associatedwith the severity of stress. The limitation to Pn was caused mainly by stomatal limitationunder moderate drought stress, and by the predominance of non-stomatal limitation undersevere stress. In this case, 40% FC water supply may be a non-stomatal limitation 5 Interspecific differences in drought tolerance of shrubs Three shrubs exhibited good performance throughout the experiment process, evenif at 20% FC treatment there were no any seedlings died, 80% FC water supply wassuitable for their establishment and growth. S. davidii minimized their water loss byreducing total leaf area and growth rate, as well as maintained higher RWC and Pncompared to the other two species under drought stress, thus they might be more tolerantto the drought stress than the other two species. On the contrary, it was found that C.polyantha and B. faberi var. microphylla had higher water loss because of their stomatalconductance and higher leaf area ratios. They reduced water loss with shedding theirleaves and changing leaf orientation under drought stress. Based on their responses, thestudied species could be categorized into two: (1) S. davidii with a tolerance mechanismin response to drought stress; (2) C. polyantha and B. faberi var. microphylla withdrought avoidance mechanism. These results indicated that slow-growing shrub speciesare better adapted to drought stress than intermediate or fast-growing species in present orpredicted drought conditions. Therefore, selecting rapid-growing species might leavethese seedlings relatively at a risk of extreme drought.

卧龙自然保护区不同海拔的中国沙棘(Hippophae rhamnoides subsp. sinensis)天然群体的遗传多样性分析

中国沙棘是一种雌雄异株、风媒传粉的灌木或乔木，在中国西南的卧龙自然保护区有广泛的分布。本研究以采集于四川卧龙自然保护区5 个海拔(1800 m、2200 m、2600 m、3000 m、3400 m)梯度的中国沙棘天然群体为材料，以ISSR 和AFLP 标记技术研究其遗传多样性水平及其遗传结构，旨在了解卧龙地区中国沙棘天然群体的遗传多样性水平以及遗传多样性在群体间、群体内以及雌雄亚群体间的分布和特征，为中国沙棘树种的遗传改良及种质资源保存提供遗传研究背景与实验依据。同时探讨ISSR、AFLP 和RAPD三种标记对中国沙棘天然群体的遗传变异水平和群体间遗传结构的评估能力和各自的优缺点。研究得出以下主要结论： 1. ISSR和AFLP分析都表明卧龙自然保护区的中国沙棘群体拥有较高的遗传变异水平(h = 0.249，HT = 0.305)。出现这种结果的主要原因可能与卧龙自然保护区多变的气候条件和生境的异质度大有关。 2. ISSR 和AFLP 都揭示出卧龙自然保护区中国沙棘群体的遗传多样性随着海拔的增加发生显著的变化，表现为中海拔群体(2200 m 和2600 m)比高海拔群体(3000 m 和3400 m)和低海拔群体(1800 m)有更高的遗传多样性的趋势。出现这种趋势的可能解释是低海拔群体处在相对高温和相对干旱的环境，高海拔群体受到低温和紫外线胁迫，而中海拔群体存在中国沙棘生长的适宜环境。 3. ISSR 和AFLP 分析都表明：卧龙自然保护区中国沙棘的遗传结构遵循分布范围广、交配系统以异交为主的木本植物的通常模式，即大多数的遗传变异存在于群体内，只有少部分的遗传变异存在于群体间。 4. 经Mantel 检测表明，卧龙自然保护区中国沙棘群体间的海拔距离和对应遗传距离之间存在显著的正相关关系，即随着垂直海拔距离的增加，群体间的遗传距离也随之增加。Mantel 检测结果以及聚类分析将卧龙自然保护区5 个不同海拔的中国沙棘群体分为低、中、高海拔群体三组的研究结果都表明，海拔很可能是限制群体间基因交流的主要因素。 5. ISSR 分析发现同一海拔的雌雄亚群体首先聚类的研究结果表明，同一海拔的雌雄亚群体在遗传上最相似。方差分析结果表明只有3.8%的总遗传变异存在于雌雄亚群体间，这可能与雌雄植株间的交配和遗传物质的混合有关。 6. ISSR、AFLP 和RAPD 分析都表明卧龙自然保护区不同海拔的中国沙棘天然群体的遗传多样性水平较高。它们的分析结果估算得到的Nei's 平均基因多样度(h)分别为0.249、0.214 和0.170。从该结果可以看出ISSR 和AFLP 比RAPD 检测到更多的遗传多态性,这很可能是不同标记检测的基因组的位点不同所致。 7. 依据对不同标记系统的比较分析，认为ISSR、AFLP 和RAPD 三种分子标记系统都能成功地用于调查卧龙自然保护区不同海拔的中国沙棘群体的遗传变异水平及遗传变异结构，提供关于中国沙棘天然群体多态性水平和遗传变异分布的有用信息。在三者中，AFLP 具有最高效能指数和标记指数，在确定种间分类关系或鉴别个体方面是一种比较理想的标记。 Hippophae rhamnoides subsp. sinensis, a dioecious and deciduous shrub species,occupies a wide range of habitats in the Wolong Nature Reserve, Southwest China. Ourpresent study investigated the pattern of genetic variation and differentiation among fivenatural populations of H. rhamnoides subsp. sinensis, occurring along an altitudinal gradientthat varied from 1,800 to 3,400 m above sea level in the Wolong Natural Reserve, by usingISSR and AFLP markers to guide its genetic improvement and germplasm conservation. And,comparative study of ISSR, AFLP and RAPD was performed to detect their capacity toestimating the level and pattern of genetic variation occurring among the five elevationpopulations of H. rhamnoides subsp. sinensis, and to discuss their application to the study onplant genetics. The results were list following: 1. The ISSR and AFLP analysis conducted for the H. rhamnoides subsp. sinensispopulations located in the Wolong Natural Reserve of China revealed the presence of highlevels of genetic variation (h = 0.249, HT = 0.305). Besides such features as relatively widedistribution, dominantly outcrossing mating system, and effective seed dispersal by small animals and birds, it is sometimes argued that hard climatic conditions and heterogeneous habitats may also contribute to high levels of diversity. 2. Genetic diversity of H. rhamnoides subsp. sinensis populations was found to varysignificantly with changing elevation, showing a trend that mid-elevation populations (2,200m and 2,600 m) were genetically more diverse than both low-elevation (1,800 m) andhigh-elevation populations (3,000 m and 3,400 m). H. rhamnoides subsp. sinensis is thoughtto be stressed by drought and high temperature at low elevations, and by low temperature athigh elevations. The high genetic variability present in the mid-elevation populations of H.rhamnoides subsp. sinensis is assumed to be related to a greater plant density in the middlealtitudinal zone, where favorable ecological conditions permit its continuous distributioncovering the zone from 2,200 m to 2,600 m above sea level. 3. The genetic structure of H. rhamnoides subsp. sinensis revealed by ISSRs andAFLPs followed the general pattern detected in woody species with widespread distributionsand outcrossing mating systems. Such plants possess more genetic diversity withinpopulations and less variation among populations than species with other combinations oftraits. 4. In the present study, Mantel tests showed positive correlations between altitudinaldistances and genetic distances among populations or subpopulations. The observedrelationship between altitude and genetic distances, and the result of the cluster analysisincluding populations or male subpopulations and classifying the groups into three altitudeclusters suggest that altitude is a major factor that restricts gene flow between populationsand subpopulations. 5. The analysis of molecular variance showed that only 3.8% of the variability residedbetween female and male subpopulations. Such a very restricted proportion of the totalmolecular variance between female and male subpopulations is due to common sexuality andmixing of genetic material between females and males. 6. The analysis based on ISSRs, AFLPs and RAPDs all revealed relatively high levelsof genetic variation among different altitudinal populations of H. rhamnoides subsp. sinensisin Wolong Natural Reserve of China. Their estimates of mean Nei’s gene diversity is equal to0.249, 0.214 and 0.170 respectively, suggesting the higher capacity of detecting geneticvariation of ISSR and AFLP than RAPD. It might be ascribed to their distinct sensitivity todifferent type of genetic variation. 7. Based on the coparative study on ISSR, AFLP and RAPD, we drew a conclusion thatthey all successfully reveal some useful information concerning the level and pattern ofgenetic vatiation occurring among different elevation populations of H. rhamnoides subsp.sinensis. AFLP is a ideal tool to taxonomic study and individual identification for theirhighest efficiency index and marker index among the three marker systems.

水分梯度上不同灌木生长对干旱河谷土壤理化和生物学性质的影响

岷江上游干旱河谷区水土流失强烈，地质灾害频繁，生态环境十分脆弱，而土壤条件恶劣（水分不足和养分缺乏）是阻碍该区植被恢复的关键因子，因此研究水分和乡土灌木生长对土壤的影响对该区的生态恢复具有指导意义。本文通过定点模拟实验，选取三种优势豆科灌木为研究对象，分别是白刺花（Sophora davidii）、小马鞍羊蹄甲（Bauhinia faberi var. microphylla）和小雀花（Campylotropics polyantha），设置5 个水分梯度，分别为100%、80%、60%、40%和20%田间持水量（FC），对栽种植物与不种植物下土壤理化性质和酶活性进行测定分析，系统比较和研究了不同水分条件和不同乡土灌木生长对干旱河谷区土壤结构、养分循环、酶活性以及微生物量的影响。主要结果如下：1. 无论生长植物与否，土壤的毛管持水量和毛管孔隙度都随着水分含量的减少而降低，最大持水量、总孔隙度和容重变化不大，相应地，土壤中的非毛管孔隙随含水量的减少而升高。各水分条件下，种植植物的毛管持水量和毛管孔隙度低于无植物生长的土壤，非毛管孔隙度相应地高于无植物土壤。土壤含水量在100%－40% FC 时，三种豆科灌木的毛管持水量和毛管孔隙度存在差异，而20% FC 条件下，三种豆科灌木土壤的物理性质基本相同。2. 水分胁迫影响土壤中养分的矿化和积累，主要表现在降低了水溶性碳和铵态氮的含量，中等程度胁迫时（60% FC）促进了有机碳和硝态氮的富集，对速效钾和有效磷没有明显作用。种植豆科灌木后各水分梯度上都增加了有机碳、铵态氮、速效钾和有效磷的积累。增加程度上三种豆科灌木间有一定差异，对于土壤有机碳总量，种植白刺花和小马鞍羊蹄甲明显高于小雀花，同样的情况还出现在铵态氮和速效钾上，但是对于有效磷，种植小雀花后的增加程度则明显高于白刺花和小马鞍羊蹄甲。种植豆科灌木不仅增加了土壤养分的相对含量，也改变了其在水分梯度上的变化趋势及其变化幅度，这种作用主要体现在碳元素和氮元素上。3. 无植物生长时脲酶活性随水分含量的减少而升高，水分胁迫对磷酸酶和过氧化氢酶的作用不显著，蔗糖酶也保持在相对较高的水平。种植植物后，蔗糖酶、磷酸酶活性与无植物时相比有较大幅度的提高，种植白刺花的脲酶活性也升高，其升高的程度在不同水分含量时不同。种植植物还降低了酶活性在水分梯度上的变幅，使之在水分梯度间的差异显著性降低。脲酶活性在指示土壤性质改变方面是较敏感的指标，其它三种酶在不同植物间的差异不明显。4. 在无植物生长时，中等程度的水分胁迫（60% FC）提高了土壤微生物量碳含量，过高或过低的土壤水分均不利于微生物碳的积累。种植小马鞍羊蹄甲后微生物量碳在水分梯度上的变化趋势与无植物生长时一致，而种植白刺花和小雀花后微生物量碳随着水分含量的减少而降低。不同种类植物的微生物量碳在水分梯度上的变化特征也不同，100% FC 条件下三种植物间没有差异，80%和60% FC 条件下小马鞍羊蹄甲显著高于白刺花和小雀花，40%和20% FC 条件下白刺花和小马鞍羊蹄甲也显著高于小雀花，说明不同种类植物随着干旱胁迫程度的加深微生物量碳的降低幅度不同，在极度干旱时，白刺花和小马鞍羊蹄甲土壤依然保持了较高的微生物活性，而小雀花土壤微生物量则明显下降。The dry valley of the upper reaches of the Minjiang River is seriously degradedmountain ecosystem. It was endangered by extremely soil lost and frequentlygeological disaster. Previous studies showed that short of water and nutrients in soilwas the principal limiting factors of vegetation restoration in this area. The typical soiland three dominant leguminous shrubs Sophora davidii, Bauhinia faberi var.microphylla and Campylotropics polyantha in upper reaches of arid Minjiang Rivervalley were considered as experimental material. Two-month old seedlings of eachspecies were exposed to five water supplies (100%, 80%, 60%, 40% and 20% waterfield capacity (FC)) in a temperature and light-controlled greenhouse. Afterthree-month water treatment, soil physiochemical variables and soil microbialactivities were determined by conventional methods. The main results showed that:1. Soil capillary capacity and capillary porosity decreased along water supplyregimes in all treatments, while saturated water capacity, total porosity and bulkdensity kept in a relatively stable level, as a result, the non-capillary porosity andcapacity increased with decrease of water supply. Compared to non-planted soil, theplant-soil systems had a higher non-capillary porosity and capacity, suggestingappropriate oxygen was present in soil to maintain the living of microorganism. Soilof three type shrub species shared the same capillary capacity and capillary porosityunder 20% FC.2. Water soluble carbon and NH4+-N decreased in response to water stress, whiletotal organic carbon and NO3--N promoted by moderate water stress and inhibited by 100% and 20% FC. Total organic carbon, NH4+-N, rapidly available K and availableP increased after the planting of leguminous shrubs in five water supply regimescompared to non-planted soil. For TOC, NH4+-N and rapidly available K, thepromotion effect was higher in S. davidii and B. faberi var. microphylla than C.polyantha planted soil, while available P displayed the opposite side. The planting ofshrubs also reduced the variance of observed traits along water supply gradients.3. Drought stress increased urease activity in non-planted soil, while insignificantdifferences were observed in phosphatase and catalase activity among five watersupply regimes. The planting of leguminous shrubs facilitated the β-glucosidase andphosphatase activity compared to the non-planted soil. It also reduced the variance ofenzyme activity along water supply gradients. Urease was more sensitive to waterstress than other three enzymes.4. Soil water content significantly affected microbial biomass carbon andCmic:Corg. S. davidii and B. faberi var. microphylla showed more drought toleranceability than C. polyantha, attributing not only to their relatively smaller variance ofmicrobial biomass carbon along soil water supply gradients, but also to the highlevel of microbial activity under severe water stress. S. davidii and B. faberi var.microphylla benefited reproduction of soil microorganism at 60%-80% FC, whilesevere drought limited it due to the competition of water and nutrients between plantand soil microorganism.

青藏高原东缘川西云杉林皆伐后灌木生长、繁殖与更新的研究

植物群落及其环境在干扰后的演替格局和过程的研究，是群落和生态系统动态研究的一个热点。选取青藏高原东缘山原区川西云杉林皆伐后，从草地过渡到灌丛的关键阶段的4 个皆伐迹地（恢复时间为8 a、10 a、16 a 和21 a），研究皆伐及自然恢复过程对林下典型灌木银露梅（Potentilla glabra）和唐古特忍冬（Lonicera tangutica）的生长与繁殖能力的影响以及灌木植物在迹地上的更新情况，分析灌木在不同生境中的适应对策和适应能力的差异,为揭示青藏高原东缘山原区迹地植被从草甸到灌丛演替的过程和特点及促进迹地演替与植被恢复进程提供理论依据和技术支撑。研究主要结论如下：1）皆伐后银露梅生长和繁殖能力显著提高，但对唐古特忍冬的影响不明显。皆伐后银露梅丛基径、高度和各部分生物量都显著增加（P < 0.05），但唐古特忍冬只有叶生物量和地下生物量增加，总生物量和其余构件生物量无显著变化。皆伐后，银露梅的结实数量、结实株数、不结实株数和结实株/不结实株比例显著增加（P < 0.05）。自然恢复过程中，银露梅和唐古特忍冬生长能力以及银露梅的结实量都表现出降低的趋势。随着迹地自然恢复时间的增加，银露梅和唐古特忍冬的基径、高度、丛叶片数和各部分生物量有减少的趋势。银露梅的结实株数增加, 但结实数量减少。2）皆伐对银露梅和唐古特忍冬生物量分配模式影响不一致。原始林和迹地中（除CT85）银露梅的生物量大小关系皆为：地下>茎>侧枝>叶。唐古特忍冬在原始林中的生物量大小关系为：茎＞地下部分＞侧枝＞叶,而皆伐后生物量的分配情况改变，生物量大小关系变为：地下部分＞茎＞侧枝＞叶。随着自然恢复时间的增加，银露梅减少了地下生物量的分配，而唐古特忍冬增加了地下生物量的分配。3）皆伐和恢复时间的增加改变了迹地物种组成，促使阳性乔、灌木在迹地上定居。4 个迹地上共出现了灌木15 种，乔木3 种，没有出现天然云杉和冷杉幼苗。随着恢复时间的增加，迹地上的灌木物种由原始林下的耐阴物种逐步发展为以针刺悬钩子（Rubus pungens）为主的阳性灌木。4）影响灌木幼苗密度和幼树密度的因子不一致。灌木幼苗密度与灌木层盖度显著负相关，与苔藓层盖度显著正相关。幼树密度与草本层盖度正相关，与苔藓层盖度、灌木层盖度和高度负相关。5）研究发现在青藏高原东缘山原区皆伐15～20 a 后，迹地仍以草本植物为主，推测皆伐后至少20 a 以上迹地才可能向灌丛阶段过渡，比高山峡谷地区的演替进程至少推迟了20 a。银露梅和唐古特忍冬在皆伐后自然恢复过程中表现出不同的生长与繁殖策略是由两个物种的生物学特性的差异引起的。银露梅比唐古特忍冬更适应迹地退化环境。促进青藏高原东缘山原林区迹地森林恢复一方面是尽量减少人为活动的破坏，另一方面，可以通过在迹地中播种适当的乡土乔、灌木种子（如白桦、银露梅）等人工措施，以加快演替进程。The succession pattern and process of plant community and their environments is a hot spotin community and ecosystem dynamic study. Four clearcuts were chosen in Rangtang(recovery time of 8 a、10 a、16 a and 21 a), which represented the key stage of thecommunity evolved from grass stage to shrub stage in the eastern margin of theQinghai-Tibetan Plateau. The growth and reproduction of the Potentilla glabra andLonicera tangutica and the natural regeneration of shrub plants in the primary Piceabalfouriana forest and 4 clearcuts were studied to explore how clear cutting andnatural recovery process affected the understory shrub species during the 21 years inthe eastern margin of the Qinghai-Tibetan Plateau. The main results were below.1) The growth and reproduction of P. glabra significantly increased after forestclear cutting.. But it was not so significant as to the L. tangutica. The organismbiomass and total biomass of P. glabra were increased obviously after clear cutting（P< 0.05）. But only leaves and underground biomass of L. tangutica increasedsignificantly after clear cutting（P < 0.05）. The number of fruit and growth of P. glabraincreased significantly after clear cutting too（P < 0.05）. The ramet height, basaldiameter , organism biomass and friut number of P. glabra and L. tangutica reducedas the increase of recovery time.2) The biomass allocation patterns varied between P. glabra and L. tangutica inthe primary forest and clearcuts. The biomass allocation of P. glabra both in primary forest and clearcuts was followed as: underground part > stem > branch > leaves.However, the biomass allocation of L. tangutica had changed after the clear cutting.The biomass allocation of L. tangutica in the primary forest was followed as: stem >underground part > branch > leaves and it was underground part > stem > branch >leave in clearcuts. The biomass allocation of P. glabra and L. tangutica varied amongclearcuts. Aboveground biomass was increased while underground biomass decreasedfor P. glabra with the increase of recovery period. However, the L. tangutica showedthe reverse changing pattern.3) Clear cutting and recovery time had changed the species composition of theclearcuts. There were 15.shruby species and 3 tree species in the four clearcuts. Nospruce and fir seedlings were found. In the early stage after clear cutting, there wereonly understory shrub species from the primary Picea balfouriana forest. The sunnyspecies, especieally Rubus pungens invaded intensly as the increase of recovery time.4) There was a significant negative relationship between density of seedlingswith shrub layer coverage and positive correlation with moss coverage. The saplingshad significantly positive correlation with herb layer coverage and negativecorrelation with moss coverage, shrub layer coverage and height.5）Comparing to studies in Miyalou, a nearby high mountain and canyon area,the secondary sucession in this subalpine plateau areas lagged at least 20 years.P. glabra and L. tangutica showed different growth and reproduction strategies toclear cutting and natural recovery , which may associated with the difference of theirbiological characters. P. glabra was more adaptive to the clear cutting than the L.tangutica. Two suggestions were probably recommended to promote the recoveryprogress in the subalpine plateau areas based on the results of this study. Limitanthropogenic disturbance, and meanwhile sow native tree and shrub seeds inclearcuts.

黄龙世界自然遗产地岷江冷杉原始林群落结构与植物多样性：生境差异性研究

黄龙世界自然遗产地岷江冷杉林(Abies faxoniana)生境类型多样，群落结构复杂，群落植物种类组成多样性丰富。揭示不同生境的生物多样性及其差异是认识生物多样性格局、形成及维持机制的前提和进行多样性保育的基础。本文采用样方法对黄龙钙化滩生境、阴坡非钙化生境及半阳坡非钙化生境的岷江冷杉原始林植物群落结构及植物多样性进行了研究。结果表明： 黄龙岷江冷杉林具有明显的复层异龄结构，垂直结构明显，乔木、灌木、草本、苔藓层次分明。共发现高等植物386 种，其中维管植物46 科103 属163 种，苔藓植38 科83 属物223 种。各层片结构及物种组成如下: (1)钙化滩生境、阴坡非钙化生境、半阳坡非钙化生境分别发现乔木18 种、13种、8 种。乔木层均可分为两个亚层，第一亚层优势种均为岷江冷杉，第二亚层主要为岷江冷杉异龄树或其它大高位芽物种。钙化滩生境第一亚层除优势种岷江冷杉外混生有巴山冷杉(Abies fargesii)、粗枝云杉(Picea asperata)以及阔叶树种白桦(Betula platyphylla)等，第二亚层主要为岷江冷杉异龄树；阴坡非钙化生境第一亚层除优势种岷江冷杉外间有巴山冷杉和白桦，第二亚层物种主要为川滇长尾槭(Acer caudatum var. prattii)；半阳坡非钙化生境第一亚层除优势种岷江冷杉外混生有巴山冷杉，第二亚层主要为岷江冷杉异龄树。依乔木层优势种的差异，钙化滩生境及半阳坡非钙化生境为岷江冷杉纯林，阴坡非钙化生境为岷江冷杉－川滇长尾槭混交林。不同生境乔木层郁闭度、乔木密度、树高结构、直径结构均存在差异。 (2)钙化滩生境发现灌木41 种，平均盖度为18.49±1.72(％)，平均高度为52.12±4.45(cm)，优势种为直穗小檗(Berberis dasystachya)；阴坡非钙化生境发现灌木30 种，平均盖度为29.33±2.56 (％)，平均高度为119.55±8.01 (cm)，优势种为箭竹 (Fargesia spathacea) 、唐古特忍冬(Lonicera tangutica) 和袋花忍冬(Lonicera saccata)；半阳坡非钙化生境发现灌木29 种，平均盖度为31.35±1.93 (％)，平均高度为107.55±4.24 (cm)，优势种为箭竹(Fargesia spathacea)。不同生境灌木层结构和物种组成多样性差异显著，钙化滩生境的灌木盖度、高度总体上较非钙化的坡地生境低, 钙化滩生境灌木以小型叶的落叶灌木为主，沟两侧非钙化的坡地生境上则发育了丰富箭竹。 (3)钙化滩生境发现草本46 种，平均盖度为7.18±0.79 (％)，平均高度为5.04±0.26(cm)，以山酢浆草(Oxalis griffithii)为优势种；阴坡非钙化生境发现草本物种71 种，平均盖度达29.04±2.31(％)，平均高度为9.08±0.52(cm)，以钝叶楼梯草(Elatostema obtusum)、山酢浆草为优势种；半阳坡非钙化生境草本物种50 种，平均盖度为以8.79±0.82(％)，平均高度为7.67±0.43 (cm)，以扇叶铁线蕨(Adiantum flabellulatum)、双花堇菜(Viola biflora)、华中蛾眉蕨(Lunathyrium shennongense)、山酢浆草为优势种。阴坡非钙化生境草本层片发育良好，多样性最为丰富，盖度和物种丰富度均显著高于钙化滩生境和半阳坡非钙化生境。 (4)钙化滩生境发现苔藓物种140 种，平均盖度达84.25±1.30 (％)，以仰叶星塔藓(Hylocomiastrum umbratum) 等大型藓类为优势种；阴坡非钙化生境发现苔藓物种115 种，平均盖度为79.29±1.64 (％)，以刺叶提灯藓(Mnium spinosum)、大羽藓(Thuidium cymbifolium)、毛尖燕尾藓(Bryhnia trichomitra)等个体较小的物种为优势种；半阳坡非钙化生境发现苔藓物种91 种，平均盖度为60.64±1.93 (％)，也以刺叶提灯藓为优势种。 (5)钙化滩生境、阴坡非钙化生境、半阳坡非钙化生境的物种数分别为234 种、221 种、175 种。乔木层的Shannon-Wiener 指数分别为0.75 ±0.12、1.87±0.12、1.78±0.07（灌木层，0.44±0.08、1.71± 0.15、2.49±0.06；草本层，0.33±0.13、1.31±0.15 、2.15±0.08； 苔藓层1.30±0.11、2.08±0.04、1.73±0.11，）；Pielou 均匀度指数分别为0.45±0.05、0.29±0.06、0.28±0.08（灌木层，0.75±0.03、0.68±0.05、0.52±0.06；草本层，0.68±0.02、0.77±0.02、0.74±0.02；苔藓层，0.40±0.03、0.63±0.02、0.52±0.03）；Simpson's 优势度指数分别为0.63±0.06、0.78±0.04、0.83±0.07（灌木层，0.21±0.03、0.28±0.05、0.45±0.06；草本层，0.25±0.02、0.12±0.01、0.17±0.01；苔藓层，0.45±0.04、0.18±0.01、0.31±0.04）。三种生境间乔木层、草本层的Sorenson 群落相似性系数较低, 灌木层、苔藓层的的Sorenson 群落相似性系数较高。 综上所述，黄龙岷江冷杉林的群落结构、植物多样性在三种生境间存在差异性，这将意味着我们在进行黄龙世界自然遗产地的森林经营管理时要较多地关注岷江冷山林群落在不同生境中的差异性。 There were multiplex habitat types, complicated community structure and abundant species composition in the Huanglong World Natural Heritage Site. Uncovering the differences of biodiversity among different habitats was a precondition to understand the distribution, formation and sustaining mechanism of the biodiversity, and the foundation of biodiversity conservation. In the present study, using plenty of quadrants, we investigated the community structure and the biodiversity of the primitive Abies faxoniana forest in different habitats (travertine bottomland, semi-sunny-slope non-calcified habitat and shady-slope non-calcified habitat) in the Huanglong World Natural Heritage Site. The main results are as follows: All the primitive Abies faxoniana forests in the three habitats were uneven-aged with obvious vertical structure including tree layer, shrub layer, herb layer and bryophyte layer. A total of 386 higher plants including 163 vascular plant species (103 generic, 46 families) and 223 bryophyte species (83 generic, 38 families) were investigated. The structure and species composition of each layer are as follows: (1) There were 18, 13 and 8 tree species in travertine bottomland, shady-slope non-calcified habitat and semi-sunny-slope non-calcified habitat, respectively. The tree layers in all habitats can be divided into two clear sub-layers. The upper tree layers were dominated by Abies faxoniana, and the lower tree layers were dominated by uneven-aged Abies faxoniana or other phanerophytes species. There were Abies fargesii , Picea asperata and Betula platyphylla besides the dominated species (Abies faxoniana) in the upper tree layer in travertine bottomland, and the lower tree layers were dominated by uneven-aged Abies faxoniana; There were Abies fargesii and Betula platyphylla besides the dominated species (Abies faxoniana) in the upper tree layer in shady-slope non-calcified habitat, and the lower tree layers were dominated by Acer caudatum var. prattii; There was Abies fargesii besides the dominated species (Abies faxoniana) in the upper tree layer semi-sunny-slope non-calcified habitat, and the lower tree layers were dominated by uneven-aged Abies faxoniana. According to composition percentage of dominate species in tree layer, both the forest in travertine bottomland and in semi-sunny-slope non-calcified habitat could be ranked as pure forest, and the forest in shady-slope non-calcified habitat could be ranked as mingled forest. There were significant differences in crown density, plant density, height structure and diameter structure among the three habitats. (2) A total of 41 shrub species (average coverage 18.49±1.72％; average height 52.12±4.45 ㎝)were found in travertine bottomland, and the dominate species was Berberis dasystachya; A total of 30 shrub species (average coverage 29.33±2.56 ％;average height 119.55±8.01 ㎝)were found in shady-slope non-calcified habitat, and the dominate species was Fargesia spathacea, Lonicera tangutica and Lonicera saccata. A total of 29 shrub species (average coverage 31.35±1.93％; average height 107.55±4.24 ㎝) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Fargesia spathacea. There were significant differences in structure and species diversity of the shrub layers among the three habitats. The coverage and height of shrub had lower value in travertine bottomland than in two non-calcified habitats. Moreover, travertine bottomland was dominated by deciduous shrub species with microphyll and non-calcified habitats developed abundant Fargesia spathacea species. (3) A total of 46 herb species (average coverage 7.18±0.79％;average height 5.04±0.26 ㎝)were found in travertine bottomland, and the dominate species was Oxalis griffithii; A total of 71 herb species (average coverage 29.04±2.31％;average height 9.08±0.52 ㎝)were found in shady-slope non-calcified habitat, and the dominate species was Elatostema obtusum and Oxalis griffithii. A total of 50 herb species (average coverage 8.79±0.82％;average height 7.67±0.43 ㎝) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Adiantum flabellulatum, Viola biflora, Lunathyrium shennongense and Oxalis griffithii. Herb layers developed well in shady-slope non-calcified habitat and had the higher species richness and coverage than travertine bottomland and semi-sunny-slope non-calcified habitat. (4) A total of 140 bryophyte species (average coverage 84.25±1.30％)were found in travertine bottomland, and the dominate species was big bryophyte species such as Hylocomiastrum umbratum and so on; A total of 115 bryophyte species (average coverage 79.29±1.64％)were found in shady-slope non-calcified habitat, and the dominate species was small bryophyte species such as Mnium spinosum, Thuidium cymbifolium, Bryhnia trichomitra and so on. A total of 91 bryophyte species (average coverage 60.64±1.93％) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Mnium spinosum. (5) There were 234, 221 and 175 plant species in travertine bottomland, shady-slope non-calcified habitat and semi-sunny-slope non-calcified habitat, respectively. Shannon-Wiener index of the tree layer was 0.75 ±0.12, 1.87±0.12 and 1.78±0.07 (the shrub layer, 0.44±0.08, 1.71± 0.15 and 2.49±0.06; the herb layer, 0.33±0.13, 1.31±0.15 and 2.15±0.08; the bryophyte layer, 1.30±0.11, 2.08±0.04 and 1.73±0.11.) for the three habitats, respectively; Pielou index of the tree layer was 0.45±0.05, 0.29±0.06 and 0.28±0.08 (the shrub layer, 0.75±0.03, 0.68±0.05 and 0.52±0.06; the herb layer, 0.68±0.02, 0.77±0.02 and 0.74±0.02; the bryophyte layer, 0.40±0.03, 0.63±0.02 and 0.52±0.03.) for the three habitats, respectively. Simpson's index of the tree layer was 0.63±0.06, 0.78±0.04 and 0.83±0.07 (the shrub layer, 0.21±0.03、0.28±0.05、0.45±0.06; the herb layer, 0.25±0.02, 0.12±0.01 and 0.17±0.01; the bryophyte layer, 0.45±0.04, 0.18±0.01 and 0.31±0.04.) for the three habitats, respectively. There were low Sorenson index both in the tree layer and in the herb layer among the three habitats, whereas, high Sorenson index occurred both in the shrub layer and in the bryophyte layer. To sum up, there were differences both in community structure and plant diversity among the three different habitats, which means that we should pay more attention to habitats heterogeneities of the primitive Abies faxoniana forest when we take action to manage the forest in the Huanglong World Natural Heritage Site.

生态恢复后的千烟洲植物群落种类组成及结构特征

Qianyanzhou(QYZ) Ecological Station established in 1983 with an area of 204 hm~2 is affiliated to the Chinese Ecosystem Research Network.Before 1982,herbs had been dominant,sparsely dotted with shrubs.After 20-year restoration of the vegetation,the vegetation showed significant changes in both forest coverage and species diversity.Forest coverage had increased to 93.3% in 1999 from 0.4% in 1982.The vegetation could be broadly classified into two groups: artificial forest,accounting for the most percent,and natural secondary forest.These two groups could be subdivided into 12 types.Based on the 2003 field work,The authors studied plant community composition and vertical structure.The results were as follows: 1) On the study plots were there about 150 species,of which 100,49,and 47 grew in arbor layer and shrub layer and herb layer,respectively.Of 12 community types,the amount of species in shrub layer was larger than that of other two layers.As to the species richness in the different community types,Liquidambar formosana community showed the highest and Imperata cylindrical var.major community the least.The amount of species in arbor layer of artificial forest was smaller than those of natural Pinus massoniana forest,but no difference in understory.2) Loropetalum chinense,Quercus fabric and Vaccinium bracteatum were dominant shrub species with a wide distribution.Three ferns Woodwardia japonica、Dryopteris atrata and Dicrannepteris dichotoma were dominant herb species.Lianas were sparse,but Milletlia reticulata were found in all forest types.3) Up to now some natural regeneration species,such as Eurya muricata、Quercus fabri、Vaccinium bracteatum、Rhus chinensis、Adinandra bockiana,had grown in the arbor layer of artificial forests.Some herb species,such as Arundinella setosa、Miscanthus floridulus、Isachne globosa、Scirpus triqueter,which were dominant ones in the herb layer before the restoration of vegetation,disappeared now.4) The vertical structure of natural Pinus massoniana community and Liquidambar formosana community showed more complex comparing with artificial forests.For the artificial forests,the conifer and broad-leaves mixed forest had a more complex structure.In both natural Pinus(massoniana) community and Liquidambar formosana community,it was dominated by individuals with height of 3～4 m,while 10～12 m in the artificial forests.

Variación del espesor cuticular foliar de cinco especies arbustivas del sur del Caldenal, Provincia de La Pampa, Argentina

p.177-182

Analyse des relations entre plasticité architecturale des buissons et prolifération de leurs populations

Réalisé en cotutelle avec l'Université Montpellier II

Systématique et biogéographie du groupe Caesalpinia (famille Leguminosae)

Parmi les lignées des Caesalpinioideae (dans la famille des Leguminosae), l’un des groupes importants au sein duquel les relations phylogénétiques demeurent nébuleuses est le « groupe Caesalpinia », un clade de plus de 205 espèces, réparties présentement entre 14 à 21 genres. La complexité taxonomique du groupe Caesalpinia provient du fait qu’on n’arrive pas à résoudre les questions de délimitations génériques de Caesalpinia sensu lato (s.l.), un regroupement de 150 espèces qui sont provisoirement classées en huit genres. Afin d’arriver à une classification générique stable, des analyses phylogénétiques de cinq loci chloroplastiques et de la région nucléaire ITS ont été effectuées sur une matrice comportant un échantillonnage taxonomique du groupe sans précédent (~84% des espèces du groupe) et couvrant la quasi-totalité de la variation morphologique et géographique du groupe Caesalpinia. Ces analyses ont permis de déterminer que plusieurs genres du groupe Caesalpinia, tels que présentement définis, sont polyphylétiques ou paraphylétiques. Nous considérons que 26 clades bien résolus représentent des genres, et une nouvelle classification générique du groupe Caesalpinia est proposée : elle inclut une clé des genres, une description des 26 genres et des espèces acceptées au sein de ces groupes. Cette nouvelle classification maintient l’inclusion de douze genres (Balsamocarpon, Cordeauxia, Guilandina, Haematoxylum, Hoffmanseggia, Lophocarpinia, Mezoneuron, Pomaria, Pterolobium, Stenodrepanum, Stuhlmannia, Zuccagnia) et en abolit deux (Stahlia et Poincianella). Elle propose aussi de réinstaurer deux genres (Biancaea et Denisophytum), de reconnaître cinq nouveaux genres (Arquita, Gelrebia, Hererolandia, Hultholia et Paubrasilia), et d’amender la description de sept genres (Caesalpinia, Cenostigma, Coulteria, Erythrostemon, Libidibia, Moullava, Tara). Les résultats indiquent qu’il y aurait possiblement aussi une 27e lignée qui correspondrait au genre Ticanto, mais un échantillonage taxonomique plus important serait nécéssaire pour éclaircir ce problème. Les espèces du groupe Caesalpinia ont une répartition pantropicale qui correspond presque parfaitement aux aires du biome succulent, mais se retrouvent aussi dans les déserts, les prairies, les savanes et les forêts tropicales humides. À l’échelle planétaire, le biome succulent consiste en une série d’habitats arides ou semi-arides hautement fragmentés et caractérisés par l’absence de feu, et abrite souvent des espèces végétales grasses, comme les Cactacées dans les néo-tropiques et les Euphorbiacées en Afrique. L’histoire biogéographique du groupe Caesalpinia a été reconstruite afin de mieux comprendre l’évolution de la flore au sein de ce biome succulent. Ce portrait biogéographique a été obtenu grâce à des analyses de datations moléculaires et des changements de taux de diversification, à une reconstruction des aires ancestrales utilisant le modèle de dispersion-extinction-cladogenèse, et à la reconstruction de l’évolution des biomes et du port des plantes sur la phylogénie du groupe Caesalpinia. Ces analyses démontrent que les disjonctions trans-continentales entre espèces sœurs qui appartiennent au même biome sont plus fréquentes que le nombre total de changements de biomes à travers la phylogénie, suggérant qu’il y a une forte conservation de niches, et qu’il est plus facile de bouger que de changer et d’évoluer au sein d’un biome différent. Par ailleurs, contrairement à nos hypothèses initiales, aucun changement de taux de diversification n’est détecté dans la phylogénie, même lorsque les espèces évoluent dans des biomes différents ou qu’il y a changement de port de la plante, et qu’elle se transforme, par exemple, en liane ou herbacée. Nous suggérons que même lorsqu’ils habitent des biomes très différents, tels que les savanes ou les forêts tropicales humides, les membres du groupe Caesalpinia se retrouvent néanmoins dans des conditions écologiques locales qui rappellent celles du biome succulent. Finalement, bien que la diversité des espèces du biome succulent ne se compare pas à celle retrouvée dans les forêts tropicales humides, ce milieu se distingue par un haut taux d’espèces endémiques, réparties dans des aires disjointes. Cette diversité spécifique est probablement sous-estimée et mérite d’être évaluée attentivement, comme en témoigne la découverte de plusieurs nouvelles espèces d’arbres et arbustes de légumineuses dans la dernière décennie. Le dernier objectif de cette thèse consiste à examiner les limites au niveau spécifique du complexe C. trichocarpa, un arbuste des Andes ayant une population disjointe au Pérou qui représente potentiellement une nouvelle espèce. Des analyses morphologiques et moléculaires sur les populations présentes à travers les Andes permettent de conclure que les populations au Pérou représentent une nouvelle espèce, qui est génétiquement distincte et comporte des caractéristiques morphologiques subtiles permettant de la distinguer des populations retrouvées en Argentine et en Bolivie. Nous décrivons cette nouvelle espèce, Arquita grandiflora, dans le cadre d’une révision taxonomique du genre Arquita, un clade de cinq espèces retrouvées exclusivement dans les vallées andines.

Root pruning as a means to encourage root growth in two ornamental shrubs, Buddleja davidii ‘Summer Beauty’ and Cistus ‘Snow Fire'

The inability of a plant to grow roots rapidly upon transplanting is one of the main factors contributing to poor establishment. In bare-rooted trees, treatments such as root pruning or application of the plant hormone auxin [e.g., indole butyric acid (IBA)] can promote root growth and aid long-term establishment. There is little information on ornamental containerised plants, however, other than the anecdotal notion that 'teasing' the roots out of the rootsoil mass before transplanting can be beneficial. In the present study we tested the ability of various root-pruning treatments and application of IBA to encourage new root and shoot growth in two shrub species, commonly produced in containers - Buddleja davidii 'Summer Beauty' and Cistus 'Snow Fire'. In a number of experiments, young plants were exposed to root manipulation (teasing, light pruning, or two types of heavy pruning) and/or treatment with IBA (at 500 or 1,000 mg l-1) before being transplanted into larger containers containing a medium of 1:1:1 (v/v/v) fine bark, sand and loam. Leaf stomatal conductance (gl) was measured 20 min, and 1, 2, 4 and 6 h after root manipulation. Net leaf CO2 assimilation (A) was measured frequently during the first week after transplanting, then at regular intervals up to 8 weeks after transplanting. Plants were harvested 8 weeks after transplanting, and root and shoot weights were measured. In both species, light root pruning alone, or in combination with 500 mg l-1 IBA, was most effective in stimulating root growth. In contrast, teasing, which is commonly used, showed no positive effect on root growth in Buddleja, and decreased new root growth in Cistus. The requirement for exogenous auxin to encourage new root growth varied between experiments and appeared to be influenced by the age and developmental stage of the plants. There were no consistent responses between root treatments and net CO2 assimilation rates, and changes in root weight were not closely correlated with changes in assimilation. The mechanisms whereby new root growth is sustained are discussed.

Adventitious root formation in anacardium occidentale L. in response to phytohormones and removal of roots

Despite advances in tissue culture techniques, propagation by leafy, softwood cuttings is the preferred, practical system for vegetative reproduction of many tree and shrub species. Species are frequently defined as 'difficult'- or 'easy-to-root' when propagated by conventional cuttings. Speed of rooting is often linked with ease of propagation, and slow-to-root species may be 'difficult' precisely because tissues deteriorate prior to the formation of adventitious roots. Even when roots form, limited development of these may impair the establishment of a cutting. In this study we used softwood cuttings of cashew (Anacardium occidentale), a species considered as 'difficult-to-root'. We aimed to test the hypothesis that speed, and extent of early rooting, is critical in determining success with this species; and that the potential to form adventitious roots will decrease with time in the propagation environment. Using two genotypes, initial rooting rates were examined in the presence or absence of exogenous auxin. In cuttings that formed adventitious roots, either entire roots or root tips were removed, to determine if further root formation/development was feasible. To investigate if subsequent root responses were linked to phytohormone action, a number of cuttings were also treated with either exogenous auxin (indole-3-butyric acid-IBA) or cytokinin (zeatin). Despite the reputation of Anacardium as being 'difficult-to-root', we found high rooting rates in two genotypes (AC 10 and CCP 1001). Removing adventitious roots from cuttings and returning them to the propagation environment, resulted in subsequent re-rooting. Indeed, individual cuttings could develop new adventitious roots on four to five separate occasions over a 9 week period. Data showed that rooting potential increased, not decreased with time in the propagation environment and that cutting viability was unaffected. Root expression was faster (8-15 days) after the removal of previous roots compared to when the cuttings were first stuck (21 days). Exposing cuttings to IBA at the time of preparation, improved initial rooting in AC 10, but not in CCP 1001. Application of IBA once roots had formed had little effect on subsequent development, but zeatin reduced root length and promoted root number and dry matter accumulation. These results challenge our hypothesis, and indicate that rooting potential remains high in Anacardium. The precise mechanisms that regulate the number of adventitious roots expressed, remain to be determined. Nevertheless, results indicate that rooting potential can be high in 'difficult-to-root' species, and suggest that providing supportive environments is the key to expressing this potential. (c) 2006 Elsevier B.V. All rights reserved.

Leaf heliotropism in Styrax camporum Pohl from the Brazilian cerrado: distinct gas exchange and leaf structure, but similar leaf temperature and water relations

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Morfoanatomia e ontogênese do pericarpo de Manihot caerulescens Pohl e M. tripartita Müll. Arg. (Euphorbiaceae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Anatomy, ultrastructure and secretion of Hibiscus pernambucensis Arruda (Malvaceae) extrafloral nectary

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Produção de sementes e comportamento germinativo de Tibouchina clavata (Pers.) Wurdack. (Melastomataceae)

In the process of riverine forest restoration, increasing emphasis has been given to the study of herbaceous and shrub species. However, for many of these we still lack basic knowledge, such as reproduction biology. Therefore, the objective of this study was to characterize the production, seed germination and reproductive capacity of Tibouchina clavata (Pers.) Wurdack., a species of wide distribution in wetlands. Fruits were collected from different mother trees located on the southern coast of São Paulo state and, after processing, were subjected to germination under constant temperatures of 25 and 30°C, and alternated temperatures of 25-30°C in the first essay, and at 15, 20, 25, 30, 35, 40°C in another essay, and at 25 °C with presence or absence of light. 10 plants were marked and, in each one, the number of fruits per plant and seed number per fruit were determined to characterize their reproductive capacity. The species germinates over a wide temperature range (15 to 35 °C), and the optimum temperature was in the range of 20 to 30 °C. Seed didn't germinate at 40 °C and could be characterized as being of absolute positive photoblastism. In general, there were over 1600 seeds per fruit and 85.000 seeds per plant. The high seed yield, coupled with good germination percentage demonstrated that this is a species with high reproductive potential.