Growth allocation between height and stem diameter in nonsuppressed reproducing Abies mariesii trees

To understand the growth patterns with respect to competition and leaf‐mass increase in reproducing trees, growth allocation between height and stem diameter was examined for nonsuppressed reproducing Abies mariesii trees in a subalpine forest in northern Honshu, Japan. The growth allocation was analyzed by dividing the relative growth rate of the stem volume into the relative contributions of height and stem‐diameter growth. During a 9‐yr period, height growth and seed‐cone production showed obvious annual variation, while stem‐diameter growth recorded moderate variation. For two of three years of seed‐cone production during the 9‐yr period, trees with larger seed‐cone production were associated with less height growth in the following year of seed‐cone production; however, there was no trend of height growth in the year of seed‐cone production. In the following year of mast seeding, trees with larger stem‐volume growth were associated with less height growth. This trend was also shown for the relationship between the cumulative stem‐volume growth during the 9‐yr period and growth allocation to height, suggesting that trees with a larger biomass increase depress the allocation of photosynthate to competition with a large expenditure for reproduction. In contrast to this, trees with a smaller biomass increase might allocate photosynthate to competition with surrounding trees. The results of this study suggest that an increase in reproductive organs during life history and annual variation in reproduction are closely associated with the growth patterns of the stem in A. mariesii trees.     

Multi-stemming strategies of Quercus glauca in an evergreen broad-leaved forest: when and where

多茎干萌生策略可赋予树木抗干扰能力。在山地生境中,树木的多茎干萌生有助于维持土壤稳定性以及森林生态系统功能的稳定性。 但对于多茎干萌生在何时(个体发育阶段偏好)、何地(生境偏好)发生,以及个体发育阶段和生境因子是否存在关联尚不了解。本文基于钱江 源国家公园古田山5 ha亚热带常绿阔叶林监测样地,研究了99个20 m × 20 m样方内常见树种青冈(Quercus glauca)萌生的发生时间与空间分 布规律。研究结果表明,青冈种群内存在3种萌生模式,大部分个体在生活史早期阶段即幼苗或者幼树阶段出现多茎干萌生。与凹凸度有关 的环境干扰可能是青冈多茎干萌生的主要驱动因素。此外,距离海拔较高的山脊越近,青冈多茎干萌生发生的越早。这一发现,强调了考虑 个体发育的重要性。因此,在研究其他森林或气候带中树种多茎干萌生策略的环境驱动因素时,要结合考虑个体发育,以期更好的理解多茎 干萌生策略及其对群落结构与功能的影响     

Canopy structure,vertical radiation profile and photosynthetic function in a Quercus ilex evergreen forest

The studied evergreen forest dominated by Quercus ilex showed a leaf area index (LAI) of 4.5, of which 61 % was accumulated within the tree layer, 30 % within the shrub layer, and 9 % within the herb layer. The leaves of all the species were ± horizontally oriented (41°), absorbing a relevant percentage of incident irradiance. The high LAI drastically modified the quality and quantity of solar radiation on the forest underground. The spectral distribution of the radiation under the forest was markedly deficient in blue and red wavelengths. The maximum absorption in these spectral bands was found in spring, when net photosynthetic rate (P _N ) was at its maximum, and in summer, when new leaves reached 90 % of their definitive structure. The vertical radiation profile showed an evident reduction of the red-far red ratio (R/FR). Radiation quality and quantity influenced leaf physiology and morphology. Clear differences in leaf size, leaf water content per area (LWC) and specific leaf area (SLA) on the vertical profile of the forest were observed. All the shrub species showed similar SLA (12.02 m2 kg-1, mean value). The ability to increase SLA whilst simultaneously reducing leaf thickness maximized the carbon economy. The high chlorophyll (Chl) content of shrub layer leaves (1.41 g kg-1, mean value) was an expression of shade adaptation. Both leaf morphology and leaf physiology expressed the phenotypic plasticity. Q. ilex, Phillyrea latifolia and Pistacia lentiscus of the forest shrub layer showed wide differences in leaf structure and function with respect to the same species developing under strong irradiance (low maquis): a 57 % mean increase of SLA and a 86 % mean decrease of PN. They showed high leaf plasticity. Leaf plasticity implies that the considered sclerophyllous species has an optimum developmental pattern achieving adaptation to environments. This revised version was published online in September 2006 with corrections to the Cover Date.     

亚热带常绿阔叶林木本植物幼树阶段抽枝展叶的权衡关系

亚热带常绿阔叶林植物幼树阶段适应林内生境并开枝散叶是其长成大树的一个重要过程, 植物一年内多次抽枝的现象及其在抽枝展叶过程中小枝伸长、枝茎增粗与叶面积的增加优先顺序及其内在驱动机制还有待进一步研究。该研究对青城山常绿阔叶林木本植物多次抽枝发生比例进行了调查, 并以茶(Camellia sinensis)、细枝柃(Eurya loquaiana)、短刺米槠(Castanopsis carlesii var. spinulosa)、润楠(Machilus nanmu)和大叶山矾(Symplocos grandis) 5种植物的幼树为研究对象, 比较分析了植物在多次抽枝中小枝和叶片生长动态及适应策略的差异。结果显示: 1)一、二次抽枝分别开始于春季(4月)和夏末(8月下旬), 小枝水平上二次抽枝率乔木小于灌木, 常绿植物小于落叶植物。2)一次抽枝小枝枝长、单叶面积, 小枝直径和叶片数量(除大叶山矾外)均高于二次抽枝, 但二次抽枝单叶面积相对生长速率均高于一次抽枝, 二次抽枝叶片比叶质量(LMA)的增长速率高于一次抽枝。3)一次抽枝小枝枝长、叶片数量、小枝直径(除细枝柃和短刺米槠外)和总叶面积(除短刺米槠外)最大相对生长速率均高于二次抽枝, 且大部分物种最大相对生长速率出现在抽枝开始的第一、二周。4)两次抽枝中, 物种先侧重于叶片的生长, 其次是小枝枝长的生长, 最后是小枝直径的增粗。单叶面积和总叶面积皆随着小枝枝长和小枝直径的增加呈显著的异速生长关系, 表明叶片的增长速度大于小枝。单叶面积与叶片数呈显著大于1的异速生长关系, 暗示单叶面积的增长速度大于叶片数的增加速度。小枝枝长与小枝直径也呈显著大于1的异速生长关系, 揭示小枝枝长的增长速度大于小枝直径。综上所述, 两次抽枝过程中, 植物枝叶的优先生长顺序反映了植物为获取更多的资源(尤其是光源)而形成特定的抽枝展叶策略; 二次抽枝单叶面积相对生长速率和LMA增长速率高于一次抽枝, 这可能与植物即将面临的昆虫取食和气温降低压力有关。因此, 了解植物抽枝策略对于理解物种生态适应机制, 揭示物种生活史过程中存在的权衡关系具有重要的理论意义。     

Tradeoff between height and relative growth rate in a dominant grass from the Serengeti ecosystem

We determined the relationship between plant height and whole-plant relative growth rate (g g^-1 day^-1) for ten genotypes of Sporobolus kentrophyllus collected from an intensively grazed site on the Serengeti Plains, Tanzania. Plants were grown for 7 weeks in a greenhouse in Syracuse, N.Y., and harvested weekly. Plants that received simulated bovine urine showed a negative relationship between plant height and growth rate, suggesting a genetic tradeoff between competitive ability if ungrazed (height) and ability to recover from grazing (growth rate). There was no height-growth rate relationship under nitrogen addition rates similar to field mineralization rates. In addition, faster-growing, shorter plants tended to have relatively higher above-ground growth rates than slower-growing, taller plants. These results suggest that natural selection has maintained a gradient of morphologies within this species ranging from short, rapidly growing genotypes adapted to intense grazing conditions to tall, slow-growing, grazer-susceptible genotypes that are superior light competitors in absence of herbivory.     

鼎湖山南亚热带常绿阔叶林荷木种群动态变化

荷木(Schima superba)是鼎湖山南亚热带常绿阔叶林群落中的优势种,为了解南亚热带常绿阔叶林的群落构建机制,该研究分析了鼎湖山南亚热带常绿阔叶林20 hm2样地中荷木种群在2005—2017年间的动态变化,并从径级和生境两个方面分析了其死亡率的分布特征。结果表明:(1)荷木种群个体数量从2005—2017年呈现出减少的态势。(2)在径级方面,荷木的死亡率和相对生长率的变化在不同径级区间呈现出不同的变化规律。径级在1～10 cm范围的荷木相对生长率大并迅速下降,死亡率上升且维持在较高水平;径级在10～50 cm范围的荷木随着径级增大,相对生长率减缓,然后下降,而死亡率不断下降;在50 cm以上的径级范围内,荷木相对生长率小、死亡率大。(3)不同生境中,2010年各个生境死亡率相差不大,2015年高海拔生境(高坡、山脊、高谷)死亡率大于低海拔生境(低坡、低谷),2017年则为低海拔生境大于高海拔生境。(4)荷木的死亡率与土壤中酸碱度呈正相关关系,而与土壤有机质、总氮、总磷、有效钾、有效氮含量呈负相关关系。     

Impacts of drought on tree mortality and growth in a mixed hardwood forest

Abstract. The tree and shrub species on a 16-ha watershed in the Coweeta Basin were sampled in 1984 and again in 1991 to determine the effects of drought on tree species composition and basal area growth. Mortality and radial growth were determined for tree species within three community types that represent a moisture gradient from moist to dry: cove-hardwoods > mixed-oak > oak-pine. Tree mortality from 1984 to 1991 was 20% and 23% in the cove-hardwoods and mixed-oak communities, respectively, compared to only 12% in the oak-pine type. With the exception of Oxydendrum arboreum and Robinia pseudoacacia, the oaks had higher percentage mortality than any other genus; Quercus velutina had 29%, 37%, and 20% mortality in the cove-hardwoods, mixed-oak, and oak-pine types, respectively; Quercus prinus had 23% mortality in the mixed-oak type; Quercus coccinea had 36% mortality in the mixed-oak type; and Quercus marilandica had 27% mortality in the oak-pine type. Mortality occurred mostly in the small-size class individuals (< 10 cm in diameter) for all species, suggesting that thinning was still an important process contributing to mortality 29 yr after clearcutting. Although growth of Liriodendron tulipifera was much higher than growth of either Quercus prinus or Quercus coccinea, growth in Liriodendron was significantly reduced by the 1985–88 drought and no growth reduction was observed for these two dominant Quercus species during the same time period.     

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...     

贵州赤水常绿阔叶林不同层次树木空间分布格局和竞争的关系

在贵州赤水桫椤国家级自然保护区的亚热带常绿阔叶林中设置0.96 hm~2的固定样地进行调查,将2299株乔木按树高分级(林木下层、中层、上层),进行空间定位并记录胸径和冠幅等个体指标。采用基于个体间距离的对数相关函数(Paircorrelation functions,PCF)和基于个体胸径、冠幅的标记点格局分析(Marked correlation functions,MCF)空间统计方法检测不同高度层内和层间的空间分布格局和潜在的竞争,结果表明:(1)样地内上层树种呈现随机分布,中下层树种在0—6 m的小尺度内呈聚集分布,在层间关系上表现为0—1 m尺度下显著的正相关关系。(2)在群落内上层个体中没有检测到显著的层内竞争,也未发现其与中、下层树种存在层间竞争关系。(3)群落中、下层树种的层内和层间竞争对两者间空间分布格局和个体大小均产生了一定影响,其相邻个体的胸径和冠幅呈现显著负相关关系。这说明在贵州赤水地区常绿阔叶林中群落中不同层次个体处于不同演替阶段,群落内层内和层间竞争对个体较大的林木上层物种的影响已经非常微弱,而主要发生在中、下层树种内,导致了个体较小的中下层更新树种在层间和层内都出现了小尺度下聚集分布格局,这解释了研究地内不同高度级别树种间竞争关系与共存策略,也验证了标记点格局分析方法在检测植物群落竞争中的敏感性。     

Trade-off between cadmium tolerance and relative growth rate in 10 grass species

Plant species exhibit great differences in heavy metal accumulation and tolerance. In this study we compared interspecific differences in responses to Cadmium (Cd) stress among 10 C₃ grass species by growing hydroponically under the conditions of different Cd treatments (0, 5, 10 and 50 μM). Responses of plant shoot dry mass to Cd treatments (resistance) were separated into avoidance (the response of shoot Cd concentration to Cd treatments) and tolerance (the response of shoot dry mass to shoot Cd concentration). The relative growth rate (RGR) and leaf structural properties of plants were measured under optimum growth condition. There were large differences in resistance, avoidance and tolerance among the species. Avoidance and tolerance were attributed independently to total Cd resistance. Resistance and tolerance were correlated negatively with RGR, leaf water content (LW), specific leaf area (SLA), leaf elongation rate (LER), and leaf length (LL), but the leaf dry matter concentration (LMDC), and nuclear DNA content showed a positive correlation with resistance and tolerance. These results indicate a trade-off between growth rate and tolerance to Cd stress. Species with higher Ca/Mg ratios showed low avoidance (r = −0.943***), suggesting uptake inhibition by Ca²⁺ to develop avoidance against Cd stress.     

Sapling biomass allocation and growth in the understory of a deciduous hardwood forest

Above- and belowground tissues of co-occurring saplings (0.1-1 m height) of Acer saccharum Marsh. (very shade tolerant), Acer rubrum L. (shade tolerant), Fraxinus americana L. (intermediate shade tolerant), and Prunus serotina Ehrh. (shade intolerant) were harvested from a forest understory to test the hypothesis that the pattern of biomass allocation varied predictably with shade-tolerance rank. The placement and length of branches along the main axis were consistent with the formation of a monolayer of foliage for the tolerant and intermediate species. Other morphological characteristics did not vary predictably with shade-tolerance rank. The maintenance of high specific leaf area (SLA; leaf area/leaf mass) and leaf area ratio (LAR; leaf area/sapling mass) is considered important for growth under extreme shade, yet these traits were not clearly related to the shade-tolerance rank of these species. Fraxinus americana, an intermediate species, had the highest LAR and growth rate in the understory, and with the exception of P. serotina, the very shade-tolerant A. saccharum had the lowest LAR. Prunus serotina maintained a large starch-rich tap root and shoot dieback was common, yielding the largest root/shoot ratio for these species. The observed allocation patterns were not similar to the long-standing expectation for the phenotypic response of juvenile trees to shade, but were consistent with three hypothetical "growth strategies" in the understory: (1) the low SLA and LAR of A. saccharum may provide a measure of defense against herbivores and pathogens and thus promote persistence in the understory, (2) the high SLA for F. americana and high LAR for F. americana and A. rubrum may enable these species to achieve high growth rates in shade, and (3) the large carbohydrate stores of P. serotina may poise this species for opportunistic growth following disturbance. The relative importance of resistance to herbivores and pathogens vs. the maintenance of high growth rates may be important in evaluating the patterns of biomass allocation in the understory.     

Stem diameter growth rates in a fire‐prone savanna correlate with photosynthetic rate and branch‐scale biomass allocation,but not specific leaf area

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (A_sat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was A_sat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. A_sat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one.     

择伐对阔叶红松林主要树种径向与纵向生长的影响

对择伐37a后的阔叶红松(Pinus koraiensis)林与原始林主要组成树种的年轮和树高数据进行分析,旨在了解择伐对其径向与纵向生长的影响。结果表明:(1)择伐显著促进了主要树种的径向生长(P0.01),但生长增加量在树种间存在显著差异(P0.01),其反应的强弱与耐荫性有关(r=-0.79,P0.01),因择伐所导致的生长加速会持续26—29a。(2)择伐显著减小了主要树种的树高-胸径比(P0.01),即同等胸径时择伐林的树木较矮,因此在计算择伐林蓄积量时应对树高进行实测。(3)径向生长率与树高、胸径之间存在显著相关性(原始林:r=0.65*,r=0.58*;择伐林:r=0.53*,r=0.48*),择伐林内每一树高级的径向生长率均高于原始林,其中树高级在10m以下的树木差异最大(0.69),说明择伐有利于林下树木的生长。择伐显著增加了DBH40cm树木的径向生长率,其中胸径在20—40cm之间的树木有较大的伐后生长率,应予以保护。     

Plants in a crowded stand regulate their height growth so as to maintain similar heights to neighbours even when they have potential advantages in height growth

Background and Aims

Although being tall is advantageous in light competition, plant height growth is often similar among dominant plants in crowded stands (height convergence). Previous theoretical studies have suggested that plants should not overtop neighbours because greater allocation to supporting tissues is necessary in taller plants, which in turn lowers leaf mass fraction and thus carbon gain. However, this model assumes that a competitor has the same potential of height growth as their neighbours, which does not necessarily account for the fact that height convergence occurs even among individuals with various biomass.

Methods

Stands of individually potted plants of Chenopodium album were established, where target plants were lifted to overtop neighbours or lowered to be overtopped. Lifted plants were expected to keep overtopping because they intercept more light without increased allocation to stems, or to regulate their height to similar levels of neighbours, saving biomass allocation to the supporting organ. Lowered plants were expected to be suppressed due to the low light availability or to increase height growth so as to have similar height to the neighbours.

Key Results

Lifted plants reduced height growth in spite of the fact that they received higher irradiance than others. Lowered plants, on the other hand, increased the rate of stem elongation despite the reduced irradiance. Consequently, lifted and lowered plants converged to the same height. In contrast to the expectation, lifted plants did not increase allocation to leaf mass despite the decreased stem length. Rather, they allocated more biomass to roots, which might contribute to improvement of mechanical stability or water status. It is suggested that decreased leaf mass fraction is not the sole cost of overtopping neighbours. Wind blowing, which may enhance transpiration and drag force, might constrain growth of overtopping plants.

Conclusions

The results show that plants in crowded stands regulate their height growth to maintain similar height to neighbours even when they have potential advantages in height growth. This might contribute to avoidance of stresses caused by wind blowing.     

Growth rates in diameter,basal area,and height of Pilgerodendron uviferum; relationship between growth index and germination

Summary Increment borer samples taken at breast height in Pilgerodendron uviferum (Cupressaceae) trees growing in two Chilean forest stands near latitude 42° south were used to study growth rates in diameter, basal area, and height. Radial average growth is 0.51 mm/year in Santa Luci 0.44 mm/year in the Piuchué stand; the correlation between diameter and age is 0.79 in Santa Lucía and 0.64 in Piuchué. Similar results were obtained in the basal area/age relationship. Both groups of trees have a low rate of growth compared to rates obtained in other studies. Maximum values for radial growth (culmination age) are reached at 110 years of age in Santa Lucia and 50 years in Piuchue. Growth in height, averaging 5.8 cm/year, appears to be similar in all trees analyzed; occasional differences can be attributed to understory position of some trees. Comparison of radial growth and germination of Pilgerodendron plants shows a synchronous relationship, suggesting an influence of climate on the pattern of forest regeneration.     

阔叶红松林不同林层和生长阶段树木生长对采伐强度的响应

采伐是调整林分结构的重要手段。不同林层的树木对采伐强度有着不同的响应方式。但以往考察采伐对树木生长的影响时多采用定性或简单定量的方法(如按树高等距)划分森林的垂直层次, 这就忽略了同一林层内不同树种间和不同发育阶段树木生长的差异。该研究在吉林蛟河天然阔叶红松(Pinus koraiensis)林内建立轻度(胸高断面积采伐强度17.3%)、中度(34.7%)、重度(51.9%)采伐以及对照(不采伐)样地, 跟踪调查采伐后自然恢复2、4、7年保留木的生长动态。根据不同树种每一个体所处的林层位置和生长发育阶段, 将保留木划分为3个组别: 林冠层树种的成熟个体(I)、林冠层树种的未成熟个体(II)以及林下层树种的全部个体(III), 比较不同恢复时期各组别树木的生长对于采伐强度的响应差异。结果表明, 第II组树木的平均胸径相对生长速率(0.033 cm·cm^-1·a^-1)显著高于第I (0.016 cm·cm^-1·a^-1)和III组(0.018 cm·cm^-1·a^-1)。总体来看, 采伐促进了大多数林冠层优势树种(第I、II组)的生长, 尤其是第II组树木的相对生长速率随采伐强度的增加而增加, 但第I组树木的相对生长速率只在重度采伐样地显著高于对照样地。然而林冠层少见种的生长速率并未受到采伐活动的显著影响。值得注意的是, 第I和II组树木生长对于采伐的响应都存在一定的时间滞后, 伐后短期内(2年)采伐样地与对照样地的生长速率没有显著差异, 而采伐对树木生长的促进效果在伐后2-4年才开始出现, 并在随后的监测期内持续存在。各组别树木的相对生长速率均随初始胸径的增大而降低, 且这种负相关关系的斜率随采伐强度增加逐渐增大, 表明随着采伐强度增加, 较小的树木个体从减弱的竞争中获益更多, 呈现出更加明显的生长释放现象。     

Plant size,environmental factors and functional traits jointly shape the stem radius growth rate in an evergreen coniferous species across ontogenetic stages

植物个体大小、环境和功能性状共同决定了常绿针叶植物个体发育阶段的径向生长率 植株个体大小、环境及功能性状对于树木生长至关重要,然而在树木个体发育过程中其对树木生长的调控机制尚不清晰。本研究以中国东北常绿针叶树种红松(Pinus koraiensis)为研究对象,选择65株胸高直径(DBH)范围为0.3–100 cm的样树,测定当年的径向生长量、环境因素(光照、土壤养分和土壤水分)以及功能性状(叶、枝和根性状)。研究结果表明,径向生长率随DBH 变化呈单峰模式,DBH为58 cm时达到峰值。结构方程模型的分析表明,当DBH < 15 cm时,植株个体大小对径向生长存在直接的负影响,但通过增加光强而间接地促进其生长;当DBH为16–58 cm和59–100 cm时,植株大小对树木径向生长分别存在直接的正影响和负影响;当DBH < 15 cm时,光照对径向生长的影响大于土壤因素,而当DBH > 15 cm时,呈相反趋势;预测树木生长的最优性状随植株个体的增大依次由比叶面积转变为木质密度及细根氮含量。因此,植株个体大小、环境和功能性状共同驱动树木的径向生长,其相对影响随树木生长而变化,且树木径向生长的限制因素随树木生长由光照转变为土壤养分。