西双版纳楝径向生长和木质部发生季节动态及其对环境因子的响应

热带森林在全球碳循环过程中起着重要的作用，研究热带树木的生长动态及其对环境因子的响应，有助于预测全球变化下热带森林固碳能力的变化。目前对热带树木径向生长以及木质部发生季节动态的研究相对较少。利用微树芯和生长环法对西双版纳热带植物园内的落叶树种楝(Melia azedarach)2021年的年内径向生长和木质部发生季节动态进行监测，结合环境因子的监测，分析了楝的径向生长过程的驱动因子。结果表明：2021年楝的扩大细胞在3月1日(年积日(DOY):60.2±1.6)开始出现，细胞壁加厚在12月11日(DOY:345.4±3.1)结束，年生长量为7.76 mm,最大生长速率为0.039 mm/d。生长环监测显示，楝在3月18日(DOY:76.6±8.6)左右开始生长，10月13日(DOY:286.4±6.8)生长结束，年径向生长量为(6.25±2.35)mm,最大生长速率为(0.056±0.02)mm/d。楝的径向生长速率与降水(r=0.77,P<0.001)、平均温度(r=0.61,P<0.05)和最低温度(r=0.67,P<0.001)均呈显著正相关，与风速(r=-0....     

热带喀斯特森林多花白头树木质部年内生长动态及其对环境因子的响应

多花白头树(Garuga floribunda var.gamblei)是西双版纳热带喀斯特森林中常见的落叶树种，容易受水分亏缺的影响。为探究热带喀斯特森林树木年内生长动态及其对环境因子的响应，该研究利用生长环和微树芯两种方法监测了多花白头树在2020—2021年的树干径向变化和木质部生长动态，并分析了树干径向变化和木质部生长与环境因子的相关性。微树芯法的监测表明，在2020年，多花白头树在3月底开始出现扩大细胞，9月底细胞壁加厚结束；在2021年，多花白头树在4月中旬开始出现扩大细胞，10月中旬细胞壁加厚结束。生长环的监测表明，2020年和2021年的径向生长开始时间均晚于扩大细胞出现时间，径向生长结束时间均早于细胞壁加厚结束时间。多花白头树在2020年和2021年的木质部生长持续时间大致相同，2020年的木质部生长量((2.87±1.46)mm)也与2021年((2.98±1.02)mm)几乎一致。多花白头树的月径向生长量、扩大细胞区域宽度和细胞壁加厚区域宽度均与降水呈显著正相关，这表明水分状况在多花白头树的木质部生长过程中发挥了重要作用；扩大细胞区域宽度还与日平均气温和日最低温显...     

树木形成层活动及其影响因素研究进展

树木生长是森林生态系统固碳的主要方式,树木生长过程受到气候与非气候因素的共同作用。树木径向生长长期定位监测是明确树木生长对气候变化响应的重要研究手段。本文对运用微树芯法的树木形成层活动及径向生长过程研究进行了总结。首先,综述了气候因素对树木形成层活动的影响: 寒冷湿润区温度决定树木生长开始和停止,干旱半干旱区水分和温度共同决定生长开始,水分决定生长停止;生长速率和持续时间共同决定生长量,最大生长速率出现在夏至前后;短期施氮并不能影响树木径向生长动态。其次,探讨了生物因素对树木径向生长过程的调控: 形成层活动开始时间因树种、树龄、竞争关系而有所差异;非结构性碳水化合物的季节动态与径向生长过程相耦合。最后,阐述了气候因素和生物因素交互作用下树木次生生长的响应机制。针对以上进展,本文提出了目前研究尚存在的问题并展望了未来的发展前景,以期为进一步的科学研究提供参考。     

热带季节雨林多花白头树年内径向生长动态及其对环境因子的响应

为了解热带地区树木的季节性生长动态和规律,在西双版纳热带季节雨林利用高精度生长仪和微树芯法对落叶树种多花白头树的径向生长季节动态进行监测。结合木质部非结构性碳水化合物和环境因子的监测,分析其形成层活动和径向季节动态的生理生态驱动因子。结果表明: 在2020年,生长仪的监测显示,多花白头树于5月底(儒略日DOY:149.3±7.2)开始生长,8月底(DOY:241.0±14.7)生长结束,年生长量为3.12 mm,最大生长速率为0.04 mm·d^-1。而微树芯法显示,扩大细胞3月9日(DOY:69.2±6.2)开始出现,9月19日(DOY:262.8±2.8)细胞加厚结束,木质部生长量为1.76 mm,最大生长速率为0.009 mm·d^-1。多花白头树径向日生长量与生长季的降水、相对湿度、日最低气温、深度为20 cm的土壤含水量和温度呈显著正相关,而与日最高气温、水汽压亏缺、最大风速和水汽压呈显著负相关。多花白头树边材淀粉含量和可溶性糖含量均在生长季开始之前保持较高水平,淀粉含量在3月底达到最低,而可溶性糖含量5月中旬达到最低,随着生长季的结束淀粉和可溶性糖的含量分别在10月中旬和12月底达到最高。     

树木年内径向生长对干旱事件的响应——以贺兰山油松为例

在贺兰山苏峪口国家森林公园,利用径向生长测量仪监测2017和2018年2个生长季内、2个海拔(2010和2330 m)油松的径向生长,研究树木径向生长对干旱的响应。结果表明: 2018年6月的干旱事件使得油松径向生长速率减慢,生长量减小;而7—8月的降水使油松的径向生长重新激活。2018年油松的径向生长主要发生在6—8月,相比2017年延长一个月。油松径向生长与气候因子的响应关系在不同海拔间没有明显的差异。生长季早期干旱对树木径向生长有抑制作用,生长季中后期降水对树木径向生长具有促进作用。该区的气候重建工作中应当充分考虑8月的气候要素对树轮宽度的影响。     

德令哈地区柏树山不同生境气候对祁连圆柏径向生长的影响

为探究德令哈地区柏树山不同生境气候对祁连圆柏（Sabina przewalskii）径向生长的影响,在该区域设置3个采样点（S1：地势较低的陡坡地,S2：坡度缓和的凹形坡地,S3：地势较高的陡坡地）,运用树木年轮学方法收集了58棵祁连圆柏共113根样芯。通过建立3个样点的祁连圆柏差值年表,分析在半干旱区不同生境内祁连圆柏径向生长对气候响应的特征以及气候因子对祁连圆柏径向生长的贡献。结果表明：不同生境的祁连圆柏径向生长对降水的响应基本一致,对温度的响应则存在显著差异,其中当年5月的降水对不同生境祁连圆柏径向生长的贡献作用最大（P<0.01）,当年6月的降水次之（P<0.05）。温度对S1和S3祁连圆柏径向生长的贡献均不显著（P>0.05）。当年9月的平均最高温对S2祁连圆柏径向生长有显著的贡献作用（P<0.05）,低温对其的贡献并不显著（P>0.05）。研究结果论证了受地形差异的影响,不同生境下的气候对树木径向生长的影响具有多样性。因此,在研究树木生长与气候的关系时,应综合考虑不同生境的特点,以选取适宜的生境进行科学研究。     

川西亚高山林线岷江冷杉和红杉对气候变化的响应

为揭示川西亚高山林线优势树种岷江冷杉（Abies fargesii var.faxoniana）和红杉（Larix potaninii）径向生长对气候变化的响应差异。通过树木年轮生态学的方法,研究了岷江冷杉和红杉径向生长与年均温的分异现象、生长衰退以及应对严重干旱事件的抵抗力和恢复力稳定性,并结合限制岷江冷杉和红杉径向生长的主要气候因素进行了分析。结果表明：（1）温度是限制林线岷江冷杉径向生长的主要气候因素,而限制红杉径向生长的主要气候因素是降水;（2）岷江冷杉的径向生长与年平均温度的变化趋势一致,红杉在20世纪90年代后径向生长呈现出下降的趋势,与年均温的升高出现分异;（3）1800-2019年间,岷江冷杉出现生长衰退2次,红杉出现生长衰退5次,相同的时间内,红杉的衰退频率高于岷江冷杉;（4）岷江冷杉对严重干旱事件的恢复力相比升温前有所增强,抵抗力相比升温前下降明显,红杉对严重干旱事件的抵抗力和恢复力相比升温前均呈现出下降的趋势。可为川西亚高山林线的植被动态变化预测和历史区域气候重建提供参考。     

论滇南西双版纳的森林植被分类

本文基于多年研究成果的总结,对西双版纳森林植被的分类、主要植被类型及其特征进行了系统归纳,并讨论了它们与世界类似热带森林植被的关系。以群落的生态外貌与结构、种类组成和生境特征相结合作为植被分类的原则和依据,可以将西双版纳的热带森林植被分类为热带雨林、热带季节性湿润林、热带季雨林和热带山地常绿阔叶林四个主要的植被型,包括有至少二十个群系。热带雨林包括热带季节雨林和热带山地(低山)雨林二个植被亚型。热带季节雨林具有与赤道低地热带雨林几乎一样的群落结构和生态外貌特征,是亚洲热带雨林的一个类型,但由于发生在季风热带北缘纬度和海拔的极限条件下,受到季节性干旱和热量不足的影响,在其林冠层中有一定比例的落叶树种存在,大高位芽植物和附生植物较逊色而藤本植物和在叶级谱上的小叶型植物更丰富,这些特征又有别于赤道低地的热带雨林。热带山地雨林是热带雨林的山地亚型,是该地区热带山地较湿润生境的一种森林类型,它在植物区系组成和生态外貌特征上类似于热带亚洲的低山雨林,隶属于广义热带雨林植被型下的低山雨林亚型。热带季节性湿润林分布在石灰岩山坡中、上部,在群落外貌上类似热带山地常绿阔叶林但在植物区系组成上与后者不同,它是石灰岩山地垂直带上的一种植被类型。热带季雨林是分布在该地区开阔河谷盆地及河岸受季风影响强烈的生境的一种热带落叶森林,是介于热带雨林与萨王纳之间的植被类型。热带山地常绿阔叶林(季风常绿阔叶林)是西双版纳的主要山地植被类型,它分布在热带季节雨林带之上偏干的山地生境。它在植物区系组成上不同于该地区的热带季节雨林,在生态外貌特征上亦不同于热带山地雨林,是发育在受地区性季风气候强烈影响的热带山地的一种森林植被类型。     

东北红豆杉雌雄植株径向生长对邻体竞争和气候因子的响应

通过树木年轮学方法, 研究了树木性别、大小、邻体竞争和气候因子(温度和降水)对东北红豆杉(Taxus cuspidata)径向生长的影响。研究表明, 性别和树木大小对东北红豆杉径向生长影响显著, 雄株的径向生长速率显著大于雌株(p=0.023)。东北红豆杉雌、雄植株径向生长速率均与邻体竞争强度显著负相关(p<0.05), 并且邻体竞争对雌株径向生长的负效应更大。树木气候学分析表明, 东北红豆杉雌、雄植株径向生长对气候因子的响应不同。雌株径向生长量与前一年8月、前一年10月平均温度显著负相关, 雄株径向生长量与月平均温度不显著相关。雌株径向生长量与当年1月的降水量显著负相关, 而雄株径向生长量与当年8月的降水显著负相关。     

阿尔泰山萨彦岭4种优势树种径向生长对气候因子的响应

气候变化深刻地影响森林树木的生长,而树种对气候变化敏感度的差异可能影响了气候变化下的森林生态系统响应。因此,研究优势树种间生长对气候变化的敏感度差异,对正确认识气候变化下林分生长动态及分布格局十分重要。基于树木年代学的方法,研究了阿尔泰山萨彦岭西伯利亚落叶松（Larix sibirica）、西伯利亚红松（Pinus sibirica）、西伯利亚冷杉（Abies sibirica）以及西伯利亚云杉（Picea obovata）4种优势树种的径向生长-气候关系。结果显示：（1）西伯利亚冷杉径向生长与上一年10-11月、当年1-9月的干旱指数、2-4月的降水显著正相关,与1月的平均温和最高温呈显著负相关关系,与当年4、6月份的水汽压正相关;（2）西伯利亚落叶松径向生长与上一年8月和当年8月的平均温、最高温以及当年8月的最低温显著负相关,而与当年6月的最低温则正相关,与8月份的水汽压显著负相关;（3）西伯利亚红松径向生长与3月降水、7月最低温、上一年10月的水汽压显著正相关;（4）西伯利亚云杉径向生长与6月平均温、最高温、水汽压正相关,与上一年10-11月、当年2-4月和9月的干旱指数正相关,同时与3、4月的降水量显著正相关。西伯利亚冷杉和西伯利亚云杉、西伯利亚云杉和西伯利亚落叶松、西伯利亚云杉和西伯利亚红松对于特定气候因子表现出相似的响应结果,与年表间相关性的结果一致。但差异也是明显的,西伯利亚冷杉和西伯利亚云杉对区域水分变化敏感,而西伯利亚落叶松和西伯利亚红松主要对区域温度变化敏感。综上所述,气候变化下,该区域优势树种对气候变化响应的差异可能导致区域林分动态和格局的改变,因此,多树种径向生长-气候关系研究有助于正确反映森林动态。研究结果可以为区域森林管理与生态保护工作提供理论依据。     

Wood density and its radial variation in six canopy tree species differing in shade-tolerance in western Thailand

Background and Aims

Wood density is a key variable for understanding life history strategies in tropical trees. Differences in wood density and its radial variation were related to the shade-tolerance of six canopy tree species in seasonally dry tropical forest in Thailand. In addition, using tree ring measurements, the influence of tree size, age and annual increment on radial density gradients was analysed.

Methods

Wood density was determined from tree cores using X-ray densitometry. X-ray films were digitized and images were measured, resulting in a continuous density profile for each sample. Mixed models were then developed to analyse differences in average wood density and in radial gradients in density among the six tree species, as well as the effects of tree age, size and annual increment on radial increases in Melia azedarach.

Key Results

Average wood density generally reflected differences in shade-tolerance, varying by nearly a factor of two. Radial gradients occurred in all species, ranging from an increase of (approx. 70%) in the shade-intolerant Melia azedarach to a decrease of approx. 13% in the shade-tolerant Neolitsea obtusifolia, but the slopes of radial gradients were generally unrelated to shade-tolerance. For Melia azedarach, radial increases were most-parsimoniously explained by log-transformed tree age and annual increment rather than by tree size.

Conclusions

The results indicate that average wood density generally reflects differences in shade-tolerance in seasonally dry tropical forests; however, inferences based on wood density alone are potentially misleading for species with complex life histories. In addition, the findings suggest that a ‘whole-tree’ view of life history and biomechanics is important for understanding patterns of radial variation in wood density. Finally, accounting for wood density gradients is likely to improve the accuracy of estimates of stem biomass and carbon in tropical trees.Key words: Radial gradients, shade-tolerance, tree biomass estimates, tree rings, tropical trees, wood density     

杉木径向生长动态及其对季节性干旱的响应

为探讨杉木径向变化的季节动态及其气候响应特征,利用径向生长仪连续2年(2016—2017年)监测了江西中部杉木的径向变化过程,分析了径向变化的日动态、季节动态规律及其与气候因子的相关性。结果表明: 杉木日径向昼夜变化呈白天收缩、夜间膨胀的格局;2017年径向生长开始时间比2016年提前一个月,但旱季持续的水分亏缺使生长季也早一个月结束;在主要生长季内(4—9月),无论湿季与旱季,径向增长量与降雨、相对湿度呈显著正相关,与光合有效辐射、饱和水汽压差呈显著负相关,而水分亏缺量的气候相关性与径向增长量相反;旱季严重缺水时土壤含水量对径向变化的影响显著增强。水分条件始终是影响杉木径向变化的关键因素,夏季干旱时可通过提高土壤含水量等有效途径促进杉木径向生长。     

Long‐term increases in intrinsic water‐use efficiency do not lead to increased stem growth in a tropical monsoon forest in western Thailand

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water‐use efficiency (iWUE). Little evidence exists, however, for the long‐term growth and gas‐exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long‐term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed‐effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance – leading to enrichment in Δ¹⁸O – than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Niño events, increased respiration, changes in allocation, or more likely, a combination of these factors.     

基于生长测量仪监测的亚热带地区马尾松多时间尺度径向变化及其与环境因子的关系

树干径向变化的多尺度研究提供了树木生长及其和环境因子关系的详细信息,有助于准确评估全球气候变化背景下森林生态系统碳汇变异。以往树干径向变化研究主要集中在温带和热带地区,且大多数研究方法基于时间分辨率较粗的树木年轮法,然而缺少亚热带地区高时间分辨率树干径向变化的研究。利用树干径向变化记录仪连续监测亚热带地区马尾松13个月的树干径向变化动态,探索不同时间尺度树干径向变化规律及与环境因子的关系。结果表明：（1）在日尺度,马尾松径向变化模式为白天收缩夜晚膨胀,秋冬季节夜晚膨胀没有春夏季明显。（2）在季节尺度,马尾松树干径向变化可分为4个时期,其中3-8月是主要生长月份,4月是累计生长量最大的月份。（3）在日尺度上,相对湿度和饱和水汽压亏缺是调节马尾松径向变化主要环境因素;在季节尺度上,土壤温度对树干径向变化的影响大于空气温度,降水量与相对湿度等水分因素对树干径向生长的促进作用在生长季中后期更为明显。研究结果有助于深入理解亚热带季风气候区树干径向变化及其对环境变化的响应,为气候变化背景下亚热带地区的植树造林设计和森林可持续管理提供依据。     

Convergence in growth responses of tropical trees to climate driven by water stress

Widely documented for temperate and cold forests in both hemispheres, variations in tree growth responses to climate along environmental gradients have rarely been investigated in the tropics. Seven tree‐ring chronologies of Centrolobium microchaete (Fabaceae) in the Cerrado tropical forests of Bolivia are used to determine the growth responses to climate along a precipitation gradient. Chronologies are distributed from the humid Guarayos forests (annual precipitation > 1600 mm) in the transition to the Amazonia to the dry‐mesic Chiquitos forests (annual precipitation < 1200 mm) in the proximity to the dry Chaco. On a large spatial scale, radial growth is positively influenced by rainfall and negatively by temperature at the end of the dry season. However, this regional pattern in climate‐tree growth relationship shows differences along the precipitation gradient. Relationships with climate are highly significant and extend over longer periods of the year in sites with low rainfall and extremely severe dry seasons. At wet sites, larger water soil capacity and endogenous forest dynamics partially mask the direct influence of climate on tree growth. Stronger similarities in tree‐growth responses to climate occur between sites in the dry Central Chiquitos and in the transition to the Guarayos forests. In contrast, the relationships show fewer similarities between sites in the humid Guarayos. We conclude that growth responses to climate in the tropics are more similar between sites with limited rainfall and severe and prolonged dry seasons. Our study points to a convergence in the patterns of growth responses of tropical trees to climate, modulated by scarce rainfall and marked seasonality. The negative impact of water deficits on tree physiological processes induces not only the documented reduction in forest species richness, but also a convergence in tree‐growth responses to climate in dry tropical forests.     

Tropical seasonal forests in monsson Asia: With emphasis on continental southeast Asia

This paper is intended to provide a brief review of the tropical seasonal forest, one type of the tropical moist forests in monsoon Asia. It will also focus on and summarise issues of current concern in relation to their depletion and global environmental issues. Tropical moist forests occur in the rainy tropical and monsoon tropical climate types. The tropical moist evergreen forest or the tropical rain forest, which account for two-thirds of the tropical moist forests are rich in biodiversity and contain valuable tropical hardwood. The tropical moist deciduous forest or the tropical seasonal forest which lie along the fringes of tropical rain forest, are less complex than the tropical rain forest and have more distinct wet and dry periods. Broadleaved deciduous trees of the genera Tectona, Shorea, and Dipterocarpus are predominantly in this forest type. Currently estimates have found that more than 17 million hectares of forest mainly tropical moist forests are being lost each year. There is a widespread recognition that agriculture and the burning of tropical moist forests contribute to global warming but to a much lesser extent than the combustion of fossil fuels and industrial activities in the developed world.     

Warming induced tree-growth decline of Toona ciliata in (sub-) tropical southwestern China

Tropical forests play an important role in the global carbon cycle and climate regulation. However, our understanding of how Asian tropical forest growth responds to climatic variations is still limited. We developed tree ring-width chronologies of Toona ciliata from 90 trees (139 cores) from two study regions in the tropical/subtropical forests in Yunnan, southwestern China. Bootstrapped correlation analysis revealed positive moisture sensitivity (precipitation, self-calibrated Palmer drought severity index, relative humidity, and soil moisture) and negative temperature sensitivity of T. ciliata, and the relationship was strongest during dry and/or dry-to-wet transition months, indicating that radial growth of T. ciliata is primarily limited by moisture availability during early growing season. Furthermore, radial growth of T. ciliata was significantly and negatively correlated with the vapor pressure deficit and potential evapotranspiration during dry and/or dry-to-wet transition months. We analyzed long-term growth trends of T. ciliata using ‘size class isolation' (SCI) and ‘generalized additive mixed models' (GAMM) approaches which remove the effects of tree size on tree growth. We detected decreasing growth trend for both approaches at both study regions, indicating that the growth decline of T. ciliata stands in southwestern China is likely due to global warming-induced moisture deficit. The growth of T. ciliata trees is likely to continually decline under projected warming and drying conditions. The observed growth declines of T. ciliata raised concerns about developing sustainable management and conservation programs for tropical/subtropical forests in China.     

Seasonal differences in leaf-level physiology give lianas a competitive advantage over trees in a tropical seasonal forest

Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO₂ assimilation per unit mass (A _mass), nitrogen concentration (N _mass), and δ¹³C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO₂ assimilation per unit area (A _area), phosphorus concentration per unit mass (P _mass), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree A _area decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana δ¹³C increased four times more than tree δ¹³C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher A _mass than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.