不同光环境下木荷幼苗树冠结构的可塑性响应

从冠形、侧枝和叶片在树冠中的空间分布等方面对天然更新木荷（Schima superba Gardn．et Champ．）幼苗的树冠结构进行了研究,认为木荷幼苗的树冠对光照条件的变化有显著的可塑性响应。随着光照水平的提高,幼苗树冠由阔、松散型向相对紧密、窄冠型发展,表明木荷幼苗对不同光照环境有较强的适应能力。木荷幼苗在强光环境下产生短枝,在适度荫庇条件下侧枝和主枝同时向上方和侧方伸长生长;在强度遮阴条件下,侧枝发生强烈的伸长生长并发生强烈的分枝行为,同时在不同自然环境条件下幼苗的叶片密度由全光、林隙到林冠下逐渐提高。随着光照水平的减弱,1级侧枝密度逐渐降低,分枝（2级侧枝和3级侧枝）强度却逐渐增大,且侧枝在树冠上的分布有向匕集中的趋势。     

桂花植冠的枝系构型分析

采用Strahler法对福建省石峰寨风景区内的不同生长发育阶段桂花(Osmanthus fragrans)的枝系构型进行了统计分析。结果表明:桂花的分枝格局参数,除总体分枝率和逐步分枝率SBR1:2无显著变化外,其余参数因发育阶段的不同,均发生不同程度的变化。枝系构型的比较分析表明:成树阶段高生长不明显,以拓展上层空间为主,幼苗阶段表现出高生长,幼树阶段则表现出从高生长向横向生长过渡的趋势。通过分析叶片配置发现:在不同枝系上,叶片均集中于植冠内一级枝和二级枝上;叶片的大小从幼苗、幼树到成树阶段逐渐增大。这些说明桂花在不同生长发育阶段,枝系构型表现出一定的可塑性,反映了不同的适应对策。     

栲树不同生长发育阶段的枝系特征分析

对浙江天童国家森林公园内常绿阔叶优势树种栲树(Castanopsis fargesii)不同发育阶段植冠内的分枝式样特征进行了统计分析。结果表明：栲树在不同发育阶段的总体分枝率和逐步分枝率有显著变化,幼苗和幼树阶段的分枝率较低,而成株阶段的分枝率较高;幼树阶段的枝条长度、枝倾角和叶倾角明显大于幼苗和成株阶段,表现为明显的高生长对策;叶片配置在不同枝系上有较大差异,叶片主要集中于植冠内一级枝和二级枝上;叶片的大小从幼苗、幼树到成株阶段逐渐增大。研究结果表明栲树在生活史的不同生长发育阶段,分枝式样表现出一定的可塑性,反映了不同的适应对策。     

不同生境栓皮栎天然更新幼苗植冠构型分析

栓皮栎存在于秦岭南坡的多种林分中,生活在不同生境中的个体往往形成不同的树冠形态和构型特征。为了说明不同生境条件下栓皮栎幼苗的植冠构型变化,采用典型抽样法,对秦岭南坡3种生境中(林冠下、林隙、林缘)的栓皮栎天然更新幼苗的侧枝、叶片特征及其空间分布进行了调查分析,结果表明:3种不同生境中栓皮栎幼苗植冠形态发生了明显的可塑性变化,(1)林冠下的幼苗明显为开阔型树冠,林隙和林缘处的幼苗树冠相对紧密;(2)幼苗的1级侧枝密度与分枝角度在3种生境下均差异显著(P0.05);从Ⅰ到Ⅳ层,林冠下幼苗的分枝角度在冠层内变化幅度不到5°,而林缘处幼苗的分枝角度变化高达40°;发生5个以上1级侧枝的概率以林冠下最大,为0.6;(3)从林缘、林隙到林冠下,幼苗的叶长、叶宽、单叶面积、叶面积指数逐渐降低,数量叶密度和比叶面积则逐渐增大,与其它两种生境相比,林冠下幼苗的叶片逐渐向树冠上层集中,且以更高序的侧枝为主要着生枝条;(4)林隙中栓皮栎幼苗的树高、地径明显优于林缘和林冠下,缩短了苗木进入主林层的时间,林隙对栓皮栎种群更新有利。在今后栓皮栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化的提供有利条件。     

不同光环境下紫椴幼树树冠结构的可塑性响应

从冠形、侧枝和叶片在树冠中的空间分布角度对天然更新紫椴幼树的树冠结构进行了论述,认为紫椴幼树树冠对光照条件的变化有显著的可塑性响应.强光通过抑制主干的生长促进了侧枝的分化,庇荫则通过抑制1级侧枝的生长促进了侧枝的再分枝.随着光照水平的降低,紫椴幼树的数量叶片密度显著降低,且叶片逐渐集中于冠上层.林冠下的紫椴幼树通过这种侧枝和叶片的分布格局,在形态上提高其对光的截获能力在适度庇荫环境中,紫椴幼树垂直生长采取演替先锋种的"避荫”对策,侧枝生长采取中等耐荫种的"掠光”对策;在弱光环境中,紫椴幼树则采取典型的忍耐适应行为.这种树冠结构的变化是提高紫椴幼树对光的截获能力的一种有益适应.     

不同光环境下胡桃楸幼苗的形态可塑性及其响应研究

为了探求不同光环境对胡桃楸幼苗生长的影响,对全光照（100%光照）、中光照（60%光照）、低光照（30%光照）3种光环境下胡桃楸幼苗的生长、形态特征、生物量分配和水分特征开展研究。结果表明：胡桃楸幼苗的形态在不同光环境下存在明显的可塑性变化,生物量分配和水分特征均表现出不同程度的差异。低光照条件下,胡桃楸幼苗生长快,株高、地径分别为全光照的1.14、1.18倍;冠幅大,是全光照的1.40倍;叶片宽大,叶长和叶宽分别是全光照的1.25和1.36倍;分枝数明显少于全光照。3种光环境下胡桃楸幼苗地上部分生物量存在显著差异,地下部分差异不显著。叶片相对含水量随着光照的降低呈现逐渐增加的趋势;全光照条件下叶片饱和持水量与叶片干鲜比明显大于遮阴条件;3种光照条件下叶绿素含量差异显著。胡桃楸幼苗为适应不同光环境,在形态和生理方面都做出适应性调整,适当的遮阴处理有利于胡桃楸幼苗生长。     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

山地落叶阔叶林优势树种米心水青冈幼苗的定居

在郁闭的林冠下、模拟林窗和林间空旷地3种光照环境中,研究了施肥和未施肥的米心水青冈幼苗生存和生长发育过程．结果表明,幼苗在林下的发育受到光照不足的严重抑制,生长在林下的幼苗比生长在林窗和空旷地幼苗死亡快,死亡率高,幼苗在郁闭林下能够生存的时间不超过12周．施肥处理的幼苗比对照的幼苗死亡快,死亡率高．幼苗死亡很可能与土壤中的病原生物有关．虽然由林下光照强度增加到模拟林窗的光照强度后,幼苗发育有明显改善,但幼苗在模拟林窗环境和空旷地的生长没有显著差异．结果说明,米心水青冈新出土的幼苗在郁闭林冠下易受伤害,更新立苗阶段需要有比较好的光照条件,肥沃土壤可能也无助于改善郁闭林冠下幼苗的定居．     

光照条件对蒙古栎幼苗生长及形态特征的影响

对生长于不同光照环境(林内和林外)下的蒙古栎幼苗的形态特征进行了比较.结果表明,光照条件对蒙古栎幼苗的生物量及其分配具有明显影响,林外幼苗单株生物量为林外的4.48倍,其中以根系生物量的差别为最大,林外为林内的6.23倍;林外与林内幼苗根冠比分别为2.70和1.11.在不同的光照条件下,蒙古栎幼苗的叶片形态表现出明显不同,林外叶片比叶面积明显低于林内,分别为139.55cm2/g 和284.94cm2/g,林内约为林外的2.04倍;一次生长叶面积林外明显低于林内,总叶面积则高于林内.二次生长使蒙古栎幼苗的叶面积有明显增加,但是经过二次生长后,幼苗的单株生物量没有明显增加,说明二次生长叶片对于蒙古栎幼苗生物量的积累没有明显贡献;光照对于蒙古栎主干及根系的形态具有明显影响,林外幼苗高(包括二次生长)明显低于林内幼苗,而地径则相反,林外幼苗根系的长度和直径则明显高于林内幼苗;林外幼苗顶芽体积明显大于林内幼苗.同时,林外幼苗的二次生长导致顶芽体积明显下降,从而对翌年的高生长量和未来干型产生影响.认为,蒙古栎幼苗通过形态上的一系列可塑性变化来适应不同的光环境以获取最大的碳收益.     

林窗对热带雨林冠层树种绒毛番龙眼幼苗生长的影响

在林窗中央、林窗边缘和林冠下3种不同光照梯度的森林生境中,研究了西双版纳季节雨林冠层树种绒毛番龙眼幼苗的早期 (种子萌发后10周内) 生长和定居后 (实生苗生长3个月以上) 的生长特点.结果表明: 绒毛番龙眼幼苗在早期生长阶段,林窗中央的株高、基径、总干质量、单株叶面积和相对生长率最大,分别为24.45 cm、3.17 mm、0.79 g、122.45 cm²和14.78×10^-3 g·d^-1.林冠下根冠比 (0.87) 高于林窗中央 (0.20) ,可能是光照和水分共同作用的结果.林窗中央较强的光照有利于定居后幼苗的生长,株高、基径、总干质量、单株叶面积、相对生长率和净同化率均在林窗中央最大,实验结束时分别达到31.48 cm、3.80 mm、2.22 g、174.52 cm²、2.29×10^-3 g·d^-1和2.54×10^-5 g·cm^-2·d^-1.幼苗死亡可能与水分胁迫密切相关,由水分胁迫引起的幼苗死亡率在林冠下最高 (26.88%),但林冠下由脊椎动物捕食引起的幼苗死亡率较低(2.93%),从而使林冠下幼苗的最终存活率最高 (70.19%).光照是影响绒毛番龙眼幼苗形态学调节的重要因素, 林窗中央不同生长阶段幼苗的比叶面积最低,但相对生长率和净同化率最大.水分胁迫和光照在幼苗定居后仍是影响幼苗生物量分配的重要因素, 林窗边缘幼苗的根冠比最高 (0.33).     

Photosynthetic responses to light in seedlings of selected Amazonian and Australian rainforest tree species

Summary Seedlings of the Caesalpinoids Hymenaea courbaril, H. parvifolia and Copaifera venezuelana, emergent trees of Amazonian rainforest canopies, and of the Araucarian conifers Agathis microstachya and A. robusta, important elements in tropical Australian rainforests, were grown at 6% (shade) and 100% full sunlight (sun) in glasshouses. All species produced more leaves in full sunlight than in shade and leaves of sun plants contained more nitrogen and less chlorophyll per unit leaf area, and had a higher specific leaf weight than leaves of shade plants. The photosynthetic response curves as a function of photon flux density for leaves of shade-grown seedlings showed lower compensation points, higher quantum yields and lower respiration rates per unit leaf area than those of sun-grown seedlings. However, except for A. robusta, photosynthetic acclimation between sun and shade was not observed; the light saturated rates of assimilation were not significantly different. Intercellular CO₂ partial pressure was similar in leaves of sun and shade-grown plants, and assimilation was limited more by intrinsic mesophyll factors than by stomata. Comparison of assimilation as a function of intercellular CO₂ partial pressure in sun- and shade-grown Agathis spp. showed a higher initial slope in leaves of sun plants, which was correlated with higher leaf nitrogen content. Assimilation was reduced at high transpiration rates and substantial photoinhibition was observed when seedlings were transferred from shade to sun. However, after transfer, newly formed leaves in A. robusta showed the same light responses as leaves of sun-grown seedlings. These observations on the limited potential for acclimation to high light in leaves of seedlings of rainforest trees are discussed in relation to regeneration following formation of gaps in the canopy.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Woodland canopy structure and the light response of juvenile Quercus lobata (Fagaceae)

I investigated competition for light between canopy plants and juvenile valley oaks (Quercus lobata Nee) in a mixed-broadleaf woodland of California's northern Coast Ranges. Canopy effects on understory light supply were separated among the overlying adult valley oak, the adult's woody understory, and neighboring trees and shrubs through a series of light sampling surveys and measurements of the number, size, and spatial distribution of neighboring plants. Light supply in the understory was primarily influenced by neighboring plants, with no detectable effect of the overlying adult valley oak. Light supply in the understory averaged 25% full sun due to a high frequency of canopy gaps and a typically open understory. Seedling response to understory light supply was investigated in an experimental sunfleck gradient (10%, 19%, and 100% full sun). Between 10% and 100% full sun, seedling growth increased by 90% and the shoot:∗∗∗root ratio changed from 1.561 to 0.607. Shade seedlings were also taller and produced fewer, larger, and thinner leaves than seedlings grown in full sun. A field survey of the spatial distribution and crown morphology of saplings and young adults found 1) the distance between young valley oaks and neighboring overstory trees to increase with neighbor size, and 2) crowns of the young oaks to be skewed away from neighbors. Although shading by the canopy was only moderate, canopy effects on understory light supply may restrict juvenile recruitment of valley oak in this woodland.     

Dependence of photosynthetic rates on leaf density thickness in deciduous woody plants grown in sun and shade

Comparisons of photosynthetic rates were made on leaves of ten species of woody dicotyledons grown in the field under full sun or under a canopy which transmitted approximately 18% of full light. Photosynthesis and dark respiration were measured and compared on various bases: area, chlorophyll, fresh weight of lamina, density thickness (fresh weight per unit area), and protein.

Light-saturated photosynthesis per unit area or unit chlorophyll was about 1.5 times greater in the sun leaves than in the shade leaves and essentially equal per unit fresh weight or unit protein. Sun leaves were thicker but the enzymes per unit fresh weight remained constant as thickness varied. Chlorophyll per unit area remained about constant; chlorophyll per unit fresh weight varied inversely with changes in leaf thickness. Thus, density thickness variation is important in photosynthetic adaptation to sun and shade. This is also shown by the relationship between light-saturated photosynthesis per unit area and density thickness.

     

Effects of canopy shade on the lipid composition of soybean leaves

The effect of canopy shade on leaf lipid composition was examined in soybeans ( Glycine max cv. Young) grown under field conditions. Expanding leaves were tagged at 50, 58 and 65 days after planting (DAP) in plots with either a high (10 plants m⁻¹ row) or low (1 plant m⁻¹ row) plant density. At 92 DAP, light conditions ranged from a pho-tosynthetic photon flux density (PPFD) of 87% of full sun with a far-red/red (735 nm/645 nm) ratio of 0.9 at upper canopy leaves to extreme shade where the PPFD was 10% of full sun with a far-red/red ratio greater than 6. Highly shaded leaves in the high plant density treatment accumulated triacylglycerol (TG) up to 25% of total leaf lipid, a 2.4-fold increase in TG on a chlorophyll basis compared to leaves in the upper canopy. Although total polar lipid content was reduced up to 50% in shaded leaves, shade had little affect on the lipid content or composition of thylakoid membranes. Shade did not affect leaf chlorophyll content. Therefore, the changes in leaf lipid composition were not related to senescence. These findings suggest that conditions of low irradiance and/or a high FR/R ratio cause a shift in carbon metabolism toward the accumulation of TG, a storage lipid. Eighteen-carbon fatty acid desaturation was also affected in highly shaded leaves where a reduction in linolenic acid (18:3) content was accompanied by a proportional increase in oleic (18:1) and linoleic (18:2) acids.     

Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees

Among 13 tropical tree species on Barro Colorado Island, species with high seedling mortality rates during the first year in shade had higher reltive growth rates (RGR) from germination to 2 months in both sun (23% full sun) and shade [2%, with and without lowered red: far red (R:FR) ratio] than shade tolerant species. Species with higher RGR in sun also had higher RGR in shade. These interspecific trends could be explained by differences in morphological traits and allocation paterns among species. Within each light regime, seedlings of shade-intolerant species had lower root: shoot ratios, higher leaf mass per unit area, and higher leaf area ratios (LAR) than shade tolerant species. In contrast, leaf gas exchange characteristics, or acclimation potential in these traits, had no relationship with seedling mortality rates in shade. In both shade tolerant and intolerant species, light saturated photosynthesis rates, dark respiration, and light compensation points were higher for sungrown seedlings than for shade-grown seedlings. Differences in R:FR ratio in shade did not affect gas exchange, allocation patterns, or growth rates of any species. Survival of young tree seedlings in shade did not depend on higher net photosynthesis or biomass accumulation rates in shade. Rather, species with higher RGR died faster in shade than species with lower RGR. This trend could be explained if survival depends on morphological characteristics likely to enhance defense against herbivores and pathogens, such as dense and tough leaves, a well-established root system, and high wood density. High construction costs for these traits, and low LAR as a consequence of these traits, should result in lower rates of whole-plant carbon gain and RGR for shade tolerant species than shade-intolerant species in shade as well as in sun.     

Ecophysiological responses of Fagus sylvatica seedlings to changing light conditions. I. Interactions between photosynthetic acclimation and photoinhibition during simulated canopy gap formation

Natural regeneration of European beech (Fagus sylvatica L.) establishes under shade, but sudden exposure to high irradiance may occur due to openings in the canopy. To elucidate ecophysiological mechanisms associated with survival of European beech seedlings, the gas exchange, chlorophyll concentrations, and chlorophyll a fluorescence parameters of two different beech populations were studied under changing light conditions. Plants were grown both in a growth chamber and at a natural site (one population) where the seedlings were raised in containers placed in understory and in simulated canopy gaps. Upon exposure to high light in the growth chamber, photosynthetic rates of shade-acclimated leaves of seedlings from both populations increased severalfold and then decreased over several days to the rates of the low-light control seedlings. High-light seedlings always had the highest photosynthetic rates. Initial fluorescence displayed a trend opposite that of photosynthesis; it increased over time, and relative fluorescence and half-time rise declined continuously until the end of experiment to very low values. Exposure to high light of shade-acclimated seedlings resulted in a shift in chlorophyll concentrations to levels intermediate between high-light and low-light seedlings. The light treatment effects were statistically greater than population effects; however, seedlings from the Abetone population were found to be more susceptible to changing light conditions than seedlings from Sicily. Reciprocal light treatments on plants growing at the natural site confirmed the results obtained in the growth chamber experiment. Overall, beech seedlings grown in the field appeared to have a fairly large acclimation potential achieved by plasticity in the photosynthetic apparatus. The lack of pronounced acclimation to high light in seedlings grown in the growth chamber was ascribed to a threshold-type relationship between the acclimation capacity and the level of damage. These observations on the limited potential for acclimation to high light in leaves of European beech seedlings which show a clear capability to exploit sunflecks, are discussed in relation to regeneration following canopy gap formation and reinforce the view of the central role of gap formation in forest dynamics. We conclude that small forest gaps (in which sunflecks play a major role) may present a favorable environment for survival and growth of beech because of their limited ability to acclimate to a sudden increase in irradiance and because of the moderate levels of light stress found in small gaps.     

Ecophysiological responses of Fagus sylvatica seedlings to changing light conditions. II. The interaction of light environment and soil fertility on seedling physiology

The survival and growth of natural beech regeneration after canopy removal is variable and little is known about ecophysiological mechanisms of these responses. Biomass, nonstructural carbohydrate levels and nitrogen concentrations were measured in an Italian population of European beech seedlings. Seedlings were container-grown in two types of soil, organic and mineral, collected at the study site. The seedlings were grown under three light treatments: under full beech canopy (understory), exposed to full sun only during midday (gap) and under full sun (clearing). Leaf gas exchange and chlorophyll a fluorescence parameters were measured and then foliar analyses were conducted for chlorophyll, phenolic and tannin levels. Biomass and allocation were significantly affected by light and soil treatments. The clearing seedlings and those in organic soil were larger than seedlings in the other light treatments or soil type. Total nonstructural carbohydrate concentrations were lower in the understory seedlings and significant differences between soil types were present in the gap and clearing seedlings. Nitrogen concentrations were higher in the understory seedlings and those growing in the organic soil compared to the other treatments. Gas exchange rates were highest in clearing and the organic soil seedlings. Gap seedlings exhibited photosynthetic acclimation that allowed them to utilize high light of midday and any sunflecks during the morning and afternoon. Relative fluorescence was significantly influenced by both light treatment and soil type, with the highest values observed in the gap seedlings. Light response curves showed decreasing apparent maximum quantum efficiency from the understory to clearing, while maximum photosynthetic rate was highest in the gap seedlings. Chlorophyll concentration was highest in understory seedlings and those growing in organic soil and higher in seedlings growing in organic than in mineral soil. Both foliar tannin and phenolic levels were highest in clearing seedlings, and only tannin concentrations were affected by soil type. Understory seedlings had the highest mortality and insect herbivory; the latter was found to be inversely related to tannin concentration. Overall, growth and photosynthesis in beech seedlings responded positively to high light associated with small canopy gaps. Organic soil increased seedling size, particularly in the gap and clearing environments. We conclude that forest gaps are favorable for photosynthesis and growth of European beech seedlings.