胡杨叶片碳水化合物及可溶性蛋白特征与叶形变化和个体发育阶段的关系

通过野外调查和室内分析,研究胡杨叶形变化及其碳水化合物及可溶性蛋白含量随个体生长发育阶段的变化规律。结果显示：（1）叶形指数随着胡杨胸径（树龄）的增加以及树冠由基向顶方向而逐渐减小,与树高和胸径均呈现极显著负相关。(2)叶片可溶性糖含量变化与个体发育阶段没有关系,在不同的发育阶段以及树冠的不同垂直空间保持在一个较高的稳定水平;叶片淀粉含量与胸径呈显著负相关,与叶形指数呈显著正相关,与叶片宽度呈极显著负相关;叶片可溶性蛋白含量与叶形指数呈显著正相关,与叶片长度呈极显著正相关。综合分析认为,胡杨叶形变化不仅存在于个体发育的不同阶段,同时存在于树冠的不同垂直空间,胡杨的叶形变化（异形叶性）是因个体发育阶段的不同而出现的生物学特征。在胡杨叶形指数随个体发育阶段、树冠由基向顶方向表现出逐渐减小的过程中,叶片淀粉含量随胸径增加、叶形指数减小和叶片宽度增加而减少;叶片可溶性蛋白含量随叶形指数和叶片长度的减小而减少。胡杨叶片淀粉、可溶性蛋白代谢可能在其叶片长度和宽度变化中具有调节作用。     

胡杨异形叶结构型性状及其相互关系

胡杨在个体生长发育过程中,叶形不断发生变化,植株上会逐年依次出现条形、披针形、卵形和阔卵形叶,且在同时具有这4种叶形的植株上,叶片在树冠中分布的顺序自上而下分别是阔卵形、卵形、披针形和条形。通过实验的方法对胡杨4种叶形叶片厚度、叶面积、比叶面积、叶片干重和干物质含量等5个结构型性状指标进行分析,研究不同叶形叶片的结构型性状与叶形(叶长/宽比值)之间的关系,以及各结构型性状指标间的相互关系。结果表明:叶片厚度、叶片干物质含量、叶面积和叶片干重与叶长/宽比值之间均呈明显负相关(P0.05);比叶面积随着叶长/宽比值的减小而逐渐减小,但二者相关性不明显,表明胡杨在个体发育过程中通过增加叶面积、叶厚度,减小比叶面积等方式来提高适应环境的能力。各结构型性状指标间相关性表现为:比叶面积与叶干物质含量和叶干重之间均呈明显负相关(P0.05),与叶厚度间呈极明显负相关(P0.01);叶片干重与叶厚度间呈明显正相关(P0.05);叶厚度与叶干物质含量之间、叶干重与叶面积之间均呈极明显正相关(P0.01),其它指标间无明显相关性。     

胡杨叶形变化与叶片养分间的关系

通过野外调查和室内分析,研究胡杨叶形随个体生长发育阶段的变化规律及其与叶片养分间的关系。结果表明:胡杨个体叶形指数(叶片长/宽)与胸径、树高和叶面积均呈极显著负相关(P0.01);不同发育阶段的胡杨均表现出叶形指数随树冠层次增高而减小的趋势,同一个体树冠不同层次的小枝也均表现为叶形指数随小枝节位数增加而减小。胡杨个体叶片全N、全P和有机C含量与树高和胸径呈正相关,全K含量与树高和胸径呈极显著负相关(P0.01);叶形指数与叶片全P含量呈显著负相关(P0.05),而与全K含量呈极显著正相关(P0.01)。说明胡杨叶形指数的变化不仅存在于个体发育的不同阶段,同时存在于同一个体树冠的不同层次以及当年生枝条的不同节位,而叶片全N、全P、全K和有机C含量的变化存在于个体发育的不同阶段。随着胸径增加,叶片叶形指数减小、叶面积增加,伴随着叶片全P、全K含量增加。     

胡杨不同发育阶段叶片光合作用及其光响应特征

胡杨(Populus euphratica)为第三纪孑遗植物, 在个体的不同发育阶段形成了独特的异形叶, 具有较强的抗旱性和耐盐性, 是额济纳绿洲生态系统的主要建群种, 具有重要的研究价值。以内蒙古额济纳旗胡杨林自然保护区胡杨为研究对象, 通过实地调查观测, 分析了胡杨幼苗期、幼树期和成熟期叶片的形态变化规律及其性状特征, 利用Li-6400XT便携式光合作用系统测定了胡杨幼苗期、幼树期和成熟期不同形态叶片的光合作用光响应过程, 并采用直角双曲线修正模型对其进行了拟合分析。结果表明: (1)在胡杨生长发育过程中, 其叶片形态性状表现出明显的变化规律。随着胡杨树龄的增长, 其叶片的长/宽比值、比叶面积和叶片厚度逐渐减小, 而叶干物质含量、叶面积和叶干重数据增大。(2)胡杨成熟期阔卵圆形叶具有较高光合活性, 通过提高P_nmax以增强光能利用率, 使其在干旱荒漠区高温、强光环境下仍能保持较高的光能转化效率与光合速率; 幼树期条形叶具有较宽的光能利用区间, 能够在弱光环境下保持较高的光合速率; 幼苗期条形叶具有较低的α与相对较高的LCP、LSP及P_nmax, 主要归因于其较高的比叶面积, 高效光捕获面积使其具有较高的光合速率; 上述结果是胡杨在不同光环境下形成的适应性反应和生理生态机制。(3)直角双曲线修正模型适于胡杨不同形态叶片光合生理参数的定量研究。     

不同展叶物候期胡杨离子分布、吸收和运输特征及其与土壤盐分关系

为明确胡杨(Populus euphratica )的离子分布、吸收和运输规律,探讨胡杨适应盐渍环境的生理机制,以不同展叶物候期（萌芽期、展叶期、叶片快速发育期、叶片成熟期）的胡杨为研究对象,测定其根、主干、老枝、幼枝和叶片中的可溶性盐离子含量,并对各离子吸收、运输和分配特征进行研究,同时结合林下土壤盐分特征,探讨胡杨离子分布与土壤盐分的关系,结果表明:随着展叶物候的变化,胡杨通过对离子吸收、运输和分布规律的差异满足生长发育的需要及对环境的适应。（1）根系选择性吸收K+、Ca2+和Mg2+的能力随展叶物候变化而呈先升后降趋势,多以叶片快速发育期为最高,恰与土壤中总盐及Na+、Cl-含量变化趋势相反。（2）离子从根系-主干及幼枝-叶片的运输能力随展叶物候的变化表现为先降后升的相反趋势,且多以叶片快速发育期为最低。（3）各器官中Na+和K+含量在展叶过程中变化不大,而在叶片成熟期有显著升高,且此期Na+以根中含量最高,K+以叶片中含量最高;叶片和根系中Ca2+、Mg2+含量随展叶物候变化整体呈升高趋势。（4）展叶物候期胡杨器官离子与土壤离子相关性表现为:萌芽期,各器官离子含量受土壤盐分影响较小,指标间相关性多不显著;展叶期,根中Na+与土壤离子成正相关,幼枝和主干K+与土壤多呈负相关;叶片快速发育期,幼枝Mg2+、根中Na+均与土壤离子呈正相关;叶片成熟期,各器官离子含量受土壤离子影响较小。     

南京地区落叶栎林主要木本植物的展叶动态研究

 植物的展叶过程是由自身遗传因子决定的,同时又受到多种生态因子的调节,反映了植物的生活史对策和群落物种多样性的维持机制。在2001和2002年的3～6月间,不定期记录了南京地区三个落叶栎(Quercus spp.)林中主要木本植物的展叶情况,包括被标记标准枝的叶数、叶的长度、宽度、叶面积、叶干重等参数。结果表明在所调查的落叶栎林中,林冠层物种的成熟叶面积和单位叶面积干重都显著大于林下层物种;最早展叶的物种为林下层物种,但林冠层展叶顺序与林下层无显著差异。叶面积越大、单位叶面积干重越小的物种展叶越晚;林冠层物种展叶较林下层快,物种成熟叶面积越大,展叶速率越大。最后对展叶顺序和展叶速度的生态学意义作了讨论。     

沈阳城市森林主要树种叶片形态生态特征与生长规律

利用统计和多元分析方法研究了沈阳城市森林10个主要树种叶片的形态生态特征和生长规律.结果表明,影响树木叶片生长发育节律的主要气象因子是≥5 ℃积温、累积日照时数和各物候期中旬平均温度.不同物候期树木所需要的气象因子有所变化,树木萌动期要求低温和寒冷指数,而在生长期需要一定的温暖指数和湿润指数.10个树种的叶片形态数量特征差异较大,按叶形指数依次为短序胡枝子＞华北丁香＞国槐＞银白杨＞红瑞木＞金银忍冬＞水腊＞水曲柳＞稠李＞黄菠萝;按叶面积依次为华北丁香＞银白杨＞黄菠萝＞稠李＞水曲柳＞红瑞木＞短序胡枝子＞金银忍冬＞国槐＞水腊.在10个树种叶长度与宽度、周长、叶面积之间相互关系中,基本属于异速生长类型,符合y=ax^k模型,但在叶长度与宽度之间关系中,除银白杨k＞1外,其它9个树种均k＜1;在叶长度与周长关系中,黄菠萝k＞1, 银白杨k≈1,呈线性关系,其它树种均k＜1;在叶长度与叶面积关系中,10个树种k＞1,而 银白杨k=2.1028,叶面积增长速率近似于叶长的2倍.建立了10个树种的叶面积最优回归估测模型,并对模型进行了估测检验.     

胡杨当年生小枝茎构型对展叶效率的影响

枝叶异速生长关系对理解荒漠植物生物量分配和生活史策略具有重要意义。该研究以小枝茎长度、茎纤细率和茎体积表征茎构型, 以叶密度(单位茎长度的叶数量)、叶面积比(单位小枝茎干质量的总叶面积)和叶茎质量比(单位小枝茎干质量的总叶干质量)表征展叶效率, 采用标准化主轴回归(SMA)方法研究胡杨(Populus euphratica)当年生小枝茎构型对展叶效率的影响及二者沿地下水埋深(GWD)梯度的权衡策略。结果显示, 胡杨当年生小枝茎直径、展叶效率和比叶面积、叶大小随GWD增加而降低, 小枝茎长度、茎纤细率及其上的叶数量则增大。小枝茎构型性状均与展叶效率呈显著负相关关系, 即随小枝茎长度、茎纤细率和茎体积的增加, 展叶效率逐渐降低, 这可能是枝叶大小、水分传导与机械支撑间的权衡结果。小枝茎构型与展叶效率的异速生长指数(斜率)随GWD增加而增大, 是由于单位小枝茎投资获得的叶面积或者叶质量降低所致, 反映出随GWD增加胡杨采取了高消耗低收益的保守型策略。胡杨应对环境压力时, 倾向于在长枝上着生数量较多的小叶, 短枝上着生数量较少的大叶, 体现出胡杨小枝的资源利用策略与枝叶大小的权衡机制。综上所述, GWD显著影响胡杨小枝茎构型-展叶效率的权衡关系, 低展叶效率是胡杨应对日益旱化荒漠环境的适应策略。     

胡杨枝、叶和花芽形态数量变化与个体发育阶段的关系

以同一立地条件下不同径级的胡杨为研究对象,通过野外调查取样和室内分析,探讨胡杨枝、叶和花芽形态数量特征随径级的变化规律及其相互间的关系。结果表明:随着径级的增加,叶形指数、枝条长度、每枝叶片数逐渐减小,叶面积、叶周长、叶柄长、枝条粗度、每枝花芽数、花芽长度和花芽宽度逐渐增加。相关分析表明,枝、叶、花芽形态数量指标均与径级存在极显著正/负相关性;花芽长度、花芽宽度、叶形指数、叶面积、叶周长、叶柄长、枝条粗度、每枝叶片数、每枝花芽数两两间均呈极显著正/负相关,枝条长度与叶形指数、叶面积、叶周长、叶柄长、枝条粗度、每枝叶片数、每枝花芽数间均呈现极显著正/负相关,说明胡杨枝、叶和花芽形态数量不仅随着个体发育阶段而逐渐变化,而且枝、叶和花芽形态数量的变化是协同进行的。方差分析显示,枝条长度、每枝叶片数在4径级开始有显著的减少,枝条粗度则与每枝花芽数、花芽长度和宽度同步从10径级开始呈现"阶梯式"的显著增加,叶形指数和叶柄长度的显著变化也出现在10径级。枝、叶和花芽形态数量表现出明显的阶段性差异与胡杨阶段转变密切相关。     

常绿和落叶阔叶物种当年生小枝茎长度和茎纤细率对展叶效率的影响

枝和叶是组成树冠的两个最主要的结构单元,枝条的结构特征及其与叶性状之间的变化关系决定了植物对光照、空间等资源的利用和适应策略。该研究以小枝茎长度和茎纤细率表征茎结构,以叶面积比(即单位小枝茎干质量的总叶面积)、叶密度(即单位茎长度的叶数量)和叶茎生物量比(即单位小枝茎干质量的总叶干质量)表征展叶效率,以浙江省清凉峰自然保护区25个常绿阔叶物种和60个落叶阔叶物种为对象,探讨小枝茎结构变化对展叶效率的影响。结果显示,展叶效率无论是用叶面积比、叶密度还是用叶茎生物量比表示,常绿和落叶物种小枝的茎长结构性状均与展叶效率呈显著的负相关关系,即随着小枝茎长度和茎纤细率的增加,展叶效率逐渐降低,可能反映了机械安全和光照限制的作用。同时,尽管茎长度-展叶效率的斜率在常绿和落叶物种间无显著差异,并且常绿物种叶密度与小枝茎结构性状之间的关系截距显著大于落叶物种,即常绿物种具有较高的出叶强度。但由于单叶面积和比叶面积显著小于落叶物种,在某一给定的小枝茎长结构下,常绿物种每单位质量的茎生物量投资获得的总叶面积和总叶质量都比落叶物种少,即常绿物种的展叶效率比落叶物种低。这可能反映了常绿物种高消耗慢收益的保守型策略。这些结果表明,小枝茎长结构对展叶效率具有显著的影响,并将随叶习性的不同而改变,与植物对环境的适应策略密切相关。     

Morphological,structural and physiological differences in heteromorphic leaves of Euphrates poplar during development stages and at crown scales

• Euphrates poplar (Populus euphratica Oliv.) has heteromorphic leaves including strip, lanceolate, ovate, and broad‐ovate leaves from base to top in the mature canopy.
• To clarify how diameter at breast height (DBH) and tree height affect the functional characteristics of all kinds of heteromorphic leaves, we measured the morphological anatomical structure and physiological indices of five crown heteromorphic leaves of P. euphratica at 2, 4, 6, 8, 10, and 12 m from the same site. We also analysed the relationships between morphological structures and physiological characteristics of heteromorphic leaves and DBH and the height of heteromorphic leaves.
• The results showed that the number of abnormalities regarding blade width, leaf area, leaf thickness, leaf mass per area, cuticle layer thickness, palisade tissue thickness, and palisade tissue/sponge tissue ratio increased with size order and sampling height gradient. Net photosynthetic rate, transpiration rate, stomatal conductance, instantaneous water use efficiency, stable delta carbon isotope ratio, proline and malondialdehyde (MDA) content increased with DBH and sampling height. By contrast, blade length, leaf shape index, and intercellular CO2 concentration decreased with the increase in path order and sampling height gradient. Although MDA content and leaf sponge thickness were not correlated with DBH or sampling height, other morphological structure and physiological parameters were significantly correlated with these variables. In addition, correlations were found among leaf morphology, anatomical structure, and physiological index parameters indicating that they changed with path order and tree height gradient.
• The differences in the morphology, anatomic structure and physiological characteristics of the heteromorphic leaves ofP. euphratica are related to ontogenesis stage and coronal position.

     

Cuticle lipids on heteromorphic leaves of Populus euphratica Oliv. growing in riparian habitats differing in available soil moisture

Populus euphratica is an important native tree found in arid regions from North Africa and South Europe to China, and is known to tolerate many forms of environmental stress, including drought. We describe cuticle waxes, cutin and cuticle permeability for the heteromorphic leaves of P. euphratica growing in two riparian habitats that differ in available soil moisture. Scanning electron microscopy revealed variation in epicuticular wax crystallization associated with leaf type and site. P. euphratica leaves are dominated by cuticular wax alkanes, primary‐alcohols and fatty acids. The major cutin monomers were 10,16‐diOH C_16:0 acids. Broad‐ovate leaves (associated with adult phase growth) produced 1.3‐ and 1.6‐fold more waxes, and 2.1‐ and 0.9‐fold more cutin monomers, than lanceolate leaves (associated with juvenile phase growth) at the wetter site and drier site, respectively. The alkane‐synthesis‐associated ECERIFERUM1 (CER1), as well as ABC transporter‐ and elongase‐associated genes, were expressed at much higher levels at the drier than wetter sites, indicating their potential function in elevating leaf cuticle lipids in the dry site conditions. Higher cuticle lipid amounts were closely associated with lower cuticle permeability (both chlorophyll efflux and water loss). Our results implicate cuticle lipids as among the xeromorphic traits associated with P. euphratica adult‐phase broad‐ovate leaves. Results here provide useful information for protecting natural populations of P. euphratica and their associated ecosystems, and shed new light on the functional interaction of cuticle and leaf heterophylly in adaptation to more arid, limited‐moisture environments.     

Identification and functional prediction of circRNAs in Populus Euphratica Oliv. heteromorphic leaves

Populus euphratica Oliv. has typical heterophylly. Linear, lanceolate, ovate and broad-ovate leaves appeared in turn from sprouting to development, to maturity. The environmental adaptabilities of P. euphraticas with different leaves were also different. To explore the role of circRNAs on the morphogenesis of P. euphratica heteromorphic leaves (P.hl) and their stress response, the expression profile of circRNAs was analyzed by strand-specific RNA sequencing for the above four kinds of heteromorphic leaves. According to ceRNA hypothesis, 18 differentially expressed cirRNAs (DECs) could influence the expression of 84 mRNAs by antagonizing 23 miRNAs in five sample-pairs. Based on the function of 84 mRNAs, these DECs participate in development process, response to stimulus, response to hormonal et al. Therefore, these circRNAs were involved in the P.hl morphogenesis and stress response by interacting with miRNAs and mRNAs. Our study complemented the genebank of P. euphratica and provided a new strategy for studying leaf development.     

胡杨异形叶光合作用对光强与CO2浓度的响应

胡杨(Populus euphratica)叶形多变, 随个体生长发育, 植株出现条形、卵形和锯齿阔卵形叶。在新疆塔里木河上游人工胡杨林内选择具有此3种叶形的成年标准株, 将枝条拉至同一高度, 通过活体测定, 比较其光合作用-光与CO₂响应及叶绿素荧光响应特征。结果表明: 胡杨异形叶光合速率对光强/CO₂浓度与电子传递速率对光强的响应曲线均可用直角双曲线修正模型来拟合, 得出的主要光合参数与实测值较吻合。胡杨卵形叶、锯齿阔卵形叶光合速率-光响应参数与生化参数及快速光响应参数与条形叶差异显著, 而光合速率-CO₂响应参数则无显著差异。胡杨异形叶CO₂饱和浓度下的最大净光合速率(P_nmax)较饱和光强下的P_nmax高, 表明胡杨强光下光合速率在很大程度上受CO₂供应和1,5-二磷酸核酮糖(RuBP)再生能力的限制。卵形叶、锯齿阔卵形叶的初始量子效率(α)、初始羧化效率(CE)、P_nmax、光合能力(A_max)与最大羧化速率(V_cmax)均显著高于条形叶; 锯齿叶光饱和点(LSP)、最大电子传递速率(ETR_max)与光呼吸速率(R_p)高于卵形叶, 条形叶光补偿点(LCP)与LSP、α、CE最低。表明荒漠干旱环境下胡杨锯齿叶最耐强光, 高R_p可能是其耗散过剩光能、保护光合机构免于强光破坏的重要途径; 卵形叶高的α、CE、磷酸丙糖利用效率(TPU)、PSII实际光化学效率(Φ_PSII)与低LCP及叶氮分配策略是其保持高光合速率的原因; 条形叶Φ_PSII、ETR、P_n低, 因其制造光合产物不足而难以满足树体生长逐渐减少并处于树冠下部。可见, 胡杨条形叶光合效率低、抗逆性差, 主要以维持生长为主; 随着树体长大, 条形叶难以适应荒漠环境来维系其生长, 出现了卵形叶; 卵形叶光合效率高, 易于快速积累光合产物而加快树体生长, 但其LSP低和耐光抑制能力弱, 逐渐被更耐强光、高温与大气干旱的锯齿叶所取代, 从而使胡杨在极端逆境下得以生存, 这是胡杨从幼苗到成年叶形变化及异形叶着生在树冠不同高度的原因。     

Phenology and intensity of phyllophage attack on Fagus sylvatica in Wytham Woods, Oxford

ABSTRACT.

• 1 Overwintering beech buds began to swell in early April and bud-break occurred in early May. Most phyllophage attacks were initiated during the 21 day period of leaf growth but additional attacks occurred up to 46 days from bud break.
• 2 An average beech tree, with a top height of 24 m and a d.b.h. of 110 cm, bore 420 345 leaves (17% in the upper canopy, 27% in the upper middle canopy, 27% in the middle canopy, 18% in the lower middle canopy and 11% in the lower canopy).
• 3 Damage by phyllophages was greatest in the lower canopy; some 75—85% of the leaves were attacked in this stratum and damage amounted to 35% of the total phyllopage damage to the trees.
• 4 The major agents of damage in the lower canopy were adult P. argentatus, larval R. fagi and lepidopteran macrophages. Adult R. fagi concentrated their feeding in the upper canopy.
• 5 Allowing for damage expansion due to leaf growth the percentages of the total leaf area actually removed, or seriously damaged, by phyllophages were calculated to be 3.43–4.00% in 1978 and 2.37–2.74% in 1979.

     

Hydraulic limitations imposed by crown placement determine final size and shape of Quercus rubra L. leaves

The canopies of large broad‐leaf trees exhibit significant heterogeneity in both micro‐environmental conditions and leaf morphology. Whether the visible differences in the size and shape of leaves from the top and bottom of the crown are determined prior to bud break or result from different patterns of leaf expansion is not known. Analysis of ontogenetic changes of both the degree of lobing and vein density in Quercus rubra demonstrates that leaves throughout the crown are identical in size and shape at the time of bud break. Morphological adaptation to the local micro‐environment takes place during the expansion phase and starts after the determination of the vascular architecture has been completed. Leaves from the bottom of the crown undergo greater expansion in the tissue close to the main veins than occurs either in the more peripheral tissue of the same leaf or anywhere in leaves from the top of the crown. This results in a water transport system that is well suited to the low evaporative rates near the bottom of the crown, but inadequate for the conditions found at the top of the tree. Acclimation of leaf form and function based upon differential expansion may be entirely driven by the local hydraulic demand during the expansion phase, resulting in leaf size and vein density being determined during development by the same hydraulic properties which will constrain the size of leaf that can be functionally supported at maturity.     

胡杨不同叶形光合特性、水分利用效率及其对加富CO2的响应

 胡杨（Populus euphratica Oliv.）叶形多变化,大致归纳为杨树叶（卵圆形叶）和柳树叶（披针形叶）两大类。在内蒙古额济纳旗胡杨林自然保护区,选择成年树同时具有卵圆形叶和披针形叶的标准株,将枝条拉至同一高度,通过活体测定,比较了其光合特征、水分利用效率及对CO2加富的响应。结果表明：在目前大气CO2浓度下,当光强为1 000 μmol·m－2·s－1时,卵圆形叶（成年树主要叶片）（A）和披针形叶（成年树下部萌条叶片）(B)的净光合速率（Pn）分别为16.40 μmol CO2·m－2·s－1和9.38 μmol CO2·m－2·s－1;水分利用效率（WUE）分别为1.52 mmol CO2·mol－1 H2O和1.18 mmol CO2·mol－1 H2O;A的光饱和点和补偿点分别为1 600 μmol·m－2·s－1和79 μmol·m－2·s－1,B的相对应值则为1 500 μmol·m m－2·s－1和168 μmol·m－2·s－1。当CO2浓度加富到450 μmol·mol－1时,A的光饱和点升高了150 μmol·m－2·s－1,光补偿点降低了36 μmol·m－2·s－1;而B的光饱和点降低了272 μmol·m－2·s－1,光补偿点则升高了32 μmol·m－2·s－1。这表明,柳树叶的光合效率较低,以维持生长为主;随着树体长大,柳树叶难以维系其生长,出现杨树叶,杨树叶更能耐大气干旱,光合效率高,通过积累光合产物,使胡杨在极端逆境下得以生存并能达到较高的生长量,这就是胡杨从幼苗到成年树叶形变化的原因。随着CO2加富,两种叶片表现出截然相反的响应,柳树叶的光合时间缩短,光能利用率减小;而杨树叶的光合时间延长,光能利用率提高。如果地下水位下降,近地层空气变干燥,或随着大气CO2浓度升高,气候变暖,柳树叶可能会逐渐减少以至消失。     

Differences in leaf phenology between juvenile and adult individuals of two tree species in a seasonally dry tropical woodland

In temperate forests, juvenile trees anticipate leaf phenology compared to adults, thus avoiding shading and herbivory. This is also expected to occur in seasonal tropical forests due to intense herbivory and shading during the rainy season; however, the anticipation of leaf phenology by juveniles in seasonal tropical forests has yet to be demonstrated. Stem‐succulent species are expected to be prone to juvenile phenological anticipation because these species are able to use water stored in their stems for leaf flushing in the dry season. We investigated this hypothesis by comparing leaf phenology (bud break, leaf expansion) of juveniles and adults of two species with contrasting wood densities in the transition between dry and rainy seasons in a tropical dry woodland. We also investigated the level of light limitation that juveniles experience in the rainy season. Both species exhibited bud break during the dry season, but only expanded their leaves with the occurrence of the first rains. In general, the stem‐succulent species had a more precocious bud break; however, anticipation by juveniles occurred only in the species with more dense wood. Canopy openness was lower than in temperate deciduous forests, but the fact that the full expansion of leaves occurred only with rainfall indicates that bud break in anticipation of canopy closure contributes only to keeping leaf photosynthetic balance from going negative, and not to higher carbon gain. The importance of anticipated budding for escaping herbivory remains an alternative explanation in need of investigation.     

Continuous and Fluctuating Leaf Phenology of Adults and Seedlings of a Shade‐tolerant Emergent Tree,Dipterocarpus sublamellatus,in Malaysia

Continuous leaf phenology, with sequential production of metamers, is usually found in pioneer species, but is rare in shade‐tolerant species. Because of the nature of ‘continuous’ growth, continuous phenology has seldom been related to meteorological factors. We studied the leaf demography of seedlings and tall trees of a shade‐tolerant emergent species, Dipterocarpus sublamellatus (Dipterocarpaceae), in Malaysia. Although leaf phenology of D. sublamellatus was classified as continuous, the rates of leaf production and leaf fall were not constant and greatly fluctuated during the census period. Seasonal patterns of leaf production and fall were not synchronized across tall trees in most combinations or between seedlings and tall trees. Mean number of leaves produced per month and its seasonal fluctuation were greater in tall trees than in seedlings. Moreover, relationships between leaf phenology and meteorological factors (monthly rainfall, net radiation, maximum and minimum temperature, and minimum relative humidity) differed among trees. This suggests that endogenous rhythms, i.e., differences in allocation of resources to growth and storage among trees, are important to fluctuations in leaf phenology. These patterns are in contrast to patterns found in various pioneer species with continuous phenology, in which rates of leaf production and fall are generally constant. Different patterns might be caused by differences in allocation to growth and storage between pioneer and shade‐tolerant species, and might be related to their regeneration strategies.