Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.     

Seasonal variation in native hydraulic conductivity between two deciduous oak species

植物抗旱性机制包括耐旱性和避旱性。耐旱性的特征主要是木质部易栓塞性和膨压损失点,而避旱性的特征则是叶片脱落和落叶性。虽然这些机制可以权衡物种连续性,但对于干旱引发的叶片脱落和持续时间的变化如何影响木质部和叶片耐受性之间的关系却知之甚少。在本研究中,我们研究两种具有不同叶片脱落叶行为的橡树(栎属)的耐旱性程度差异。我们预测了Quercus deserticola 在旱季结束时落叶(晚落叶)而面临更大的空化风险,导致其比Q. laeta更耐旱,对水分的利用也更加保守。Q. laeta 只在旱季中期很短的时间内落叶(短落叶)。本研究在墨西哥中部进行,两种橡树各具有单个种群,它们之间相距1.58公里,海拔差为191米。Q. deserticola (晚落叶)多出现在下坡,而Q. laeta (短落叶)多出现在沿坡度海拔较高的地方。我们评估了样地内茎干的水分传导率的季节差异(雨季和旱季),并检测了两个物种间木质部对空化的脆弱性、叶片水分利用和叶片膨松损失点的差异。两种橡树在耐旱性状方面没有显著差异,主要包括木质部易栓塞性、叶片膨松损失点和气孔导度。然而,两种植物在旱季都有不同的表现：短落叶植物对木质部功能的负面影响要比晚落叶植物低。综上所述,两种橡树在植物生理性能方面的季节变化取决于冠层叶面积的减少。     

Coloration and phenology manifest nutrient variability in senesced leaves of 46 temperate deciduous woody species

颜色和物候表明46种温带落叶木本植物衰老叶片的养分变异 不同共生植物的叶片养分含量差异显著,反映了不同的叶片养分利用策略。然而,衰老叶片养分的种间变异及其驱动因素尚不清楚。本研究旨在探讨衰老叶片养分的种间变异及其驱动因素。我们在中国东北的帽儿山森林生态系统研究站测定了46种共存温带落叶木本植物新鲜落叶的碳、氮、磷浓度。 采用随机森林模型量化10个生物因素(菌根类型、固氮类型、生长形态、耐阴性、叶片质地、变色程度、变色类型、叶片变色峰期、落叶峰期和落叶末期)的相对重要性。研究结果表明,落叶氮浓度种间变化为4倍,磷浓度变化达9倍。较高的氮和磷平均浓度(15.38和1.24 mg g⁻¹)表明该森林氮和磷限制较弱。功能群仅对特定养分及其比值有显著影响。磷浓度、氮磷比与外生菌根树种的落叶高峰日和落叶结束日呈负相关。颜色鲜艳的叶片(红色＞棕色＞黄色＞黄绿色＞绿色)倾向于比绿色叶片氮和磷浓度更低而碳氮比和碳磷比较高。随机森林模型表明,秋季叶变色和落叶物候贡献了80%的种间变异解释量。这些结果增加了我们对温带森林木本植物营养策略之衰老叶片养分变异性的理解。     

不同光环境下亚热带常绿阔叶树种和落叶阔叶树种幼苗的叶形态和光合生理特征

由于地球环境的演变,亚热带常绿阔叶林中常有落叶阔叶成分的存在,但从生态角度,其存在的机理尚不清楚.通过比较研究常绿阔叶优势树种大头茶与落叶阔叶树种枫香幼苗的叶形态和光合生理特征对不同光环境(旷地、林窗、林下)的响应,尝试解释落叶成分遗留的基础.结果表明:①枫香具有较小的比叶重(LMA)和较高的光合氮利用效率(PNUE),在高光环境中,并未受到光抑制,具有较高的光合可塑性,尤其在林窗,表现出较大头茶高3倍的光合能力(Pnmax),保证了其在较短的生长季节积累更多的光合产物,增强与常绿树种的竞争力;②常绿阔叶树种大头茶具有较大的LMA,在旷地受到严重的光抑制,将更多的氮用于化学防御中.在林窗和林下具有相对高的Pnmax,生长幅度较广,但在林下将更多的氮投入到比生长更为重要的生存消耗中.而枫香在林下响应表现为Rubisco活性降低,光合受阻;③两树种在林窗均表现出较高的光合适应性,具有最大的光合能力(Pnmax)及合理的氮在光合机构中的分配系数,说明林窗是它们更新的最佳环境.但枫香的光合作用更依赖于光照,而大头茶则更依赖于CO2浓度;④总之,落叶阔叶树种以其较高的形态可塑性和对高光的光合生理可塑性能够在常绿阔叶林种生存、生长,并成为常绿阔叶林中固有成分的主要原因之一.     

林冠结构和地形对亚热带常绿落叶阔叶林林下幼苗物种多样性和功能多样性的影响

森林结构和地形是森林生态系统最明显的特点,也是影响林下幼苗存活和物种多样性的关键因子。该研究采用半球面摄影方法提取八大公山生长监测样地（共1.2 hm²）林冠结构参数,通过调查地表层木本植物幼苗的组成和多度,获取常见植物幼苗叶片功能性状,结合详细的地形信息,利用空间同步自回归模型探究林冠结构变量及地形因子对幼苗物种多样性及功能多样性的影响。结果表明：（1）八大公山亚热带山地常绿落叶阔叶林林冠结构复杂度较高,最大林冠高的平均值达到19.94 m,叶面积指数、平均叶倾角和林冠覆盖度分别为2.94、30.88°和0.87;（2）林冠结构变量和地形因子能够解释32.6%~48.4%的林下幼苗物种多样性指数变异和28.5%~70.2%的功能多样性变异,但地形因子对幼苗物种多样性的影响很小;（3）预测在亚热带常绿落叶阔叶林高海拔的山坡上,有较低的叶面积指数和平均叶倾角群落有较高的幼苗物种多样性;而在低海拔山脊上,较低的叶面积指数和平均叶倾角群落林下幼苗层有较高的功能多样性。此结果对科研人员和林业工作者开展野外森林更新情况评估和样方调查将有所帮助。     

Diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in India

BACKGROUND AND AIMS: In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species. METHODS: Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles. KEY RESULTS: Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3-4 months) in dry-season and winter-flowering species, 6-9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized. CONCLUSIONS: The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.     

哀牢山中山湿性常绿阔叶林木径向生长季节动态及其对气候因子的响应

研究采用树木生长环在哀牢山中山湿性常绿阔叶林持续9年(2009—2017年)监测了2个常绿树种(厚皮香,Ternstroemia gymnanthera;南亚枇杷,Eriobotrya bengalensis)和2个落叶树种(西桦,Betula alnoides;珍珠花,Lyonia ovalifolia)的树干月生长量,采用逻辑斯蒂生长模型(Logistic model)模拟树木径向生长量和物候参数,并分析了年、季尺度上径向生长与主要气候因子的关系。结果表明:1)4个树种年平均生长量为6.3 mm,落叶树种年平均生长量(10.6 mm/a)显著高于常绿树种(3.0 mm/a);2)雨季(5—10月)是哀牢山中山湿性常绿阔叶林树木生长的主要时期,4个树种雨季平均生长量为5.9 mm,占全年总生长量的93%,其中落叶树种雨季生长量占全年的96%,而常绿树种雨季生长量占全年的86%;3)常绿树种生长季长度为169天,长于落叶树种(137天),而落叶树种最大生长速率(0.14 mm/d)显著高于常绿树种(0.03 mm/d),最大径向生长速率能很好地预测树种年生长量;4)低温、雾日和光合有效...     

Effects of size,neighbors, and site condition on tree growth in a subtropical evergreen and deciduous broad‐leaved mixed forest,China

Successful growth of a tree is the result of combined effects of biotic and abiotic factors. It is important to understand how biotic and abiotic factors affect changes in forest structure and dynamics under environmental fluctuations. In this study, we explored the effects of initial size [diameter at breast height (DBH)], neighborhood competition, and site condition on tree growth, based on a 3‐year monitoring of tree growth rate in a permanent plot (120 × 80 m) of montane Fagus engleriana–Cyclobalanopsis multiervis mixed forest on Mt. Shennongjia, China. We measured DBH increments every 6 months from October 2011 to October 2014 by field‐made dendrometers and calculated the mean annual growth rate over the 3 years for each individual tree. We also measured and calculated twelve soil properties and five topographic variables for 384 grids of 5 × 5 m. We defined two distance‐dependent neighborhood competition indices with and without considerations of phylogenetic relatedness between trees and tested for significant differences in growth rates among functional groups. On average, trees in this mixed montane forest grew 0.07 cm year^?1 in DBH. Deciduous, canopy, and early‐successional species grew faster than evergreen, small‐statured, and late‐successional species, respectively. Growth rates increased with initial DBH, but were not significantly related to neighborhood competition and site condition for overall trees. Phylogenetic relatedness between trees did not influence the neighborhood competition. Different factors were found to influence tree growth rates of different functional groups: Initial DBH was the dominant factor for all tree groups; neighborhood competition within 5 m radius decreased growth rates of evergreen trees; and site condition tended to be more related to growth rates of fast‐growing trees (deciduous, canopy, pioneer, and early‐successional species) than the slow‐growing trees (evergreen, understory, and late‐successional species).     

Shoot structure and dynamics of saplings and canopies of three deciduous broad-leaved trees of a coppice forest in central Japan

Three deciduous broad-leaved trees, Quercus serrata, Castanea crenata and Carpinus laxiflora, were the main constituents of a coppice forest in central Japan. The shoot elongation and leaf emergence modes of both saplings and the canopy of the three species were investigated. The shoot elongation modes of Q. serrata and C. crenata were the same in saplings and the upper layer of the canopy. The second shoots of these two species were formed after the first shoots were elongated. C. laxiflora was different between saplings and the upper layer of the canopy. In saplings, only the first shoots took a long time to elongate. In the upper canopy layer, higher order shoots were formed in the same way as in the other two species. In the lower layer of the canopy, all three species showed the same shoot elongation mode, in which only the first shoot and its duration of elongation was short. Leaf longevity, individual leaf area, leaf mass per unit leaf area and the stem mass per unit stem length of C. laxiflora were significantly shorter or significantly smaller than those of Q. serrata and C. crenata. The length of the stem per unit leaf area of C. laxiflora was three times that of Q. serrata and five times that of C. crenata. The elongation growth of C. laxiflora was highly efficient as it occurred with a small leaf area. The shoot dynamics and the shoot structure of C. laxiflora are more suitable for elongation growth than in Q. serrata and C. crenata. Furthermore, the shoot structures of the three species were compared and ecological characteristics of the three species are discussed. Received: 29 September 1998 / Accepted: 17 September 1999     

Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species

Photosynthetic nitrogen use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) is one of the most important factors for the interspecific variation in photosynthetic capacity. PNUE was analysed in two evergreen and two deciduous species of the genus Quercus. PNUE was lower in evergreen than in deciduous species, which was primarily ascribed to a smaller fraction of nitrogen allocated to the photosynthetic apparatus in evergreen species. Leaf nitrogen was further analysed into proteins in the water‐soluble, the detergent‐soluble, and the detergent‐insoluble fractions. It was assumed that the detergent‐insoluble protein represented the cell wall proteins. The fraction of nitrogen allocated to the detergent‐insoluble protein was greater in evergreen than in deciduous leaves. Thus the smaller allocation of nitrogen to the photosynthetic apparatus in evergreen species was associated with the greater allocation to cell walls. Across species, the fraction of nitrogen in detergent‐insoluble proteins was positively correlated with leaf mass per area, whereas that in the photosynthetic proteins was negatively correlated. There may be a trade‐off in nitrogen partitioning between components pertaining to productivity (photosynthetic proteins) and those pertaining to persistence (structural proteins). This trade‐off may result in the convergence of leaf traits, where species with a longer leaf life‐span have a greater leaf mass per area, lower photosynthetic capacity, and lower PNUE regardless of life form, phyllogeny, and biome.     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

表征亚热带常绿林光合作用季节变化特征的多种植被指数

利用遥感方法可以在区域尺度反演地表植被的光合生理状况和生产力变化,但亚热带常绿林冠层结构季节变化较小,传统的光谱植被指数对植被光合作用难以准确捕捉。利用2014—2015年中国科学院广东省鼎湖山森林生态试验站多角度自动光谱观测系统的光谱反射数据,分别反演传统冠层结构型植被指数(NDVI)、光合生理生化型植被指数(CCI)和叶绿素荧光型植被指数(NDFI_(685)和NDFI_(760)),并利用不同类型植被指数的组合,构建多元线性回归模型。结果表明:亚热带常绿针阔混交林三种类型植被指数均与GPP的动态变化有显著的相关性,其中,NDVI是表征GPP较优的植被指数(R~2=0.60,P0.01),其次为CCI(R~2=0.55,P0.01),而NDFI能够作为辅助指数,有效提高NDVI(R~2=0.68,P0.001)和CCI(R~2=0.67,P0.001)表征GPP的程度。多个植被指数参与构建的多元回归模型能够有效提高亚热带地区常绿林GPP季节动态变化的拟合精度,提升遥感精确评估亚热带森林生产力的能力。     

Hydraulic architecture of deciduous and evergreen dry rainforest tree species from north-eastern Australia

Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (K_s) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of K_h and leaf specific conductivity (K_L) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.     

喀斯特常绿落叶阔叶林木本植物功能性状变异及其适应策略

叶片功能性状能反映植物对环境的高度适应能力和复杂生境下的自我调控能力,同时也能反映植物的基本特征和对资源的有效利用率。以木论国家级自然保护区喀斯特常绿落叶阔叶林144种优势木本植物为研究对象,测定其叶厚(LT)、叶面积(LA)、比叶面积(SLA)、叶长宽比(L/W)、叶组织密度(LTD)叶片形态性状和12种叶元素性状特征及变异程度,并探讨植物对喀斯特生境的适应策略。结果表明：17个性状变异程度不同,其中叶面积变异系数最大,达到133.31%,叶片碳变异系数最小,为7.73%,叶元素变异程度普遍高于叶形态性状变异程度。不同叶习性物种间叶厚、比叶面积、叶长宽比、叶氮含量性状差异达到显著水平。部分叶性状呈显著相关,得到一系列最佳功能性状组合,体现植物对喀斯特地区特殊生境的适应性。沿着性状贡献率较高的PC1轴,能够定义出叶经济谱,大部分常绿植物采取经济保守策略,而大部分落叶植物则聚集在资源获取的一侧。相较于邻近非喀斯特地区,喀斯特地区植物有较小的LA、SLA。这些结果体现了喀斯特地区植物叶片形成的不同叶性状特征,以及分布于经济谱两端的常绿和落叶植物的不同资源获取策略,揭示了植物对生境的适应策略...     

Comparing growth phenology of co-occurring deciduous and evergreen conifers exposed to drought

Plant phenological events are influenced by climate factors such as temperature and rainfall. To evaluate phenological responses to water availability in a Spring Heath-Pine wood (Erico-Pinetum typicum), the focus of this study was to determine intra-annual dynamics of apical and lateral growth of co-occurring early successional Larix decidua and Pinus sylvestris and late successional Picea abies exposed to drought. The effect of reduced plant water availability on growth phenology was investigated by conducting a rainfall exclusion experiment. Timing of key phenological dates (onset, maximum rate, end, duration) of growth processes were compared among species at the rain-sheltered and control plot during 2011 and 2012. Shoot and needle elongation were monitored on lateral branches in the canopy at c. 16 m height and radial growth was recorded by automatic dendrometers at c. 1.3 m height of >120 yr old trees. Different sequences in aboveground growth phenology were detected among the three species under the same growing conditions. While onset of radial growth in April through early May was considerably preceded by onset of needle growth in Larix decidua (5–6 weeks) and shoot growth in Pinus sylvestris (c. 3 weeks), it occurred quite simultaneously with onset of shoot growth in Picea abies. Low water availability had a minor impact on onset of aboveground growth, which is related to utilization of stored water, but caused premature cessation of aboveground growth. At the control plot mean growing season length was 130 days in Pinus sylvestris, 95 days in Larix decidua and 73 days in Picea abies, supporting the hypothesis that early successional species are resource expenders, while late successional species are more efficient in utilizing resources and develop safer life strategies. High synchronicity found in culmination of radial growth in late spring (mid-May through early June) prior to occurrence of more favourable environmental conditions in summer might indicate sink competition for carbohydrates to belowground organs. This is supported by completion of apical growth in mid June in all species, except for needle growth of Pinus sylvestris which lasted until early August. Phenological observations of conifers exposed to drought revealed that tree water status early during the growing season determines total annual aboveground growth and, besides temperature, species-specific endogenous and/or environmental factors (most likely photoperiod and/or different threshold temperatures) are involved in controlling apical and lateral growth resumption after winter dormancy.     

Ultrasound acoustic emissions from dehydrating leaves of deciduous and evergreen trees

While drying, detached leaves produced ultrasound acoustic emissions (UAE) comparable to emissions from stem and twig wood. Experiments on Ilex aquifolium L. showed that the main source of these signals was cavitation in the veins, to which conduits and fibres probably both contributed. Regions of the leaf blade with abundant mesophyll and only small veins emitted few signals. More signals were counted on the adaxial side of the midrib than on the abaxial one and on the proximal third than on the distal one, in accordance with the anatomical structure. Sound attenuation was pronounced. Eight species were compared with respect to cavitation behaviour, field water relations and pressure–volume curves, and the data showed differences in cumulative number of events and resistance of leaves to cavitation. Data were generally in good agreement with anatomical structure and habitat preferences. The number of signals per conduit counted on cross-sections was in some leaves much higher than unity, which suggests short xylem elements or an acoustic activity of cells other than conduits. There was no correlation between cavitation threshold or cumulative number of signals and the degree of sclerophylly; unexpectedly, there was a correlation between the cumulative number of signals at a water potential of -1.3 MPa and the bulk modulus of elasticity.     

The structure and phenology of a moist deciduous forest in the Central Indian Highlands

The vegetation structure and phenology of a 74.5 ha block of moist deeiduous forest in Kanha Tiger Reserve, Madhya Pradesh, India were investigated during primatological fieldwork. Within a grid of 0.25 ha quadrats all 9935 trees (woody plants > 2 m tall), of 63 species, were enumerated and their girth measured. The forest was dominated by the dipterocarp sal (Shorea robusta). Most species were rare and showed clumped distributions within the sea of sal whilst the dispersion of five large canopy trees could not be distinguished from random.The phenology of 215 trees, of 61 species, was monitored over 14 months using an index of phytophase abundance. Leaf renewal was highly synchronous between and within species, mostly occurring between February and June. One species was evergreen, 5 were semi-evergreen and 55 were deciduous. Flowering generally occurred between leaf fall and flush whilst fruiting peaked late in the hot weather and early monsoon.