遮荫对江南油杉幼苗生长和叶绿素荧光参数的影响北大核心CSCD

为探究有利于江南油杉(Keteleeria fortunei var.cyclolepis)幼苗生长发育的最适光照条件,该研究以1.5年生江南油杉幼苗为研究对象,对其进行为期2年不同光照强度(即全光照和遮荫率40%、60%、80%)处理,分析其形态生长、叶绿素含量、叶绿素荧光参数、生物量积累及分配等对不同光环境的适应机制。结果表明：(1)随遮荫程度的增强,江南油杉幼苗的地径生长量、叶片厚度逐渐减小,株高生长量、叶面积和比叶面积逐渐增大,叶片干质量呈先升后降的趋势;(2)遮荫造成叶片的叶绿素含量、初始荧光(Fo)、PSⅡ潜在活性(Fv/Fo)、PSⅡ最大光化学量子产量(Fv/Fm)、PSⅡ实际光化学量子产量[Y(Ⅱ)]、电子传递速率(ETR)均呈现显著增加的趋势;(3)遮荫处理下幼苗的叶、茎、根生物量和总生物量总体表现出随着光照强度的降低而减小的变化趋势,幼苗根生物量比和根冠比均减小,而叶生物量比、茎生物量比以及光合组织与非光合组织生物量比均增大。研究发现,全光照下江南油杉幼苗生长发育受到了抑制,遮荫显著影响江南油杉苗期生长及叶绿素荧光特性,适度遮荫能提高幼苗叶片的光捕获能力、光能利用效率,增强幼苗光合能力,促进生长;江南油杉幼苗在遮荫率为60%的环境下受光胁迫较小,其生长、叶片表观特征、光合能力以及生物量积累与分配等表现最优。     

不同生境朝鲜淫羊藿生长与光合特征

选择朝鲜淫羊藿野外生存的林缘、林窗和林下3种生境,研究朝鲜淫羊藿形态特征、光合特征、叶绿素含量、生物量积累及分配对不同光环境的响应。结果表明,朝鲜淫羊藿株高、茎径、总叶面积等形态指标随光照强度减弱明显减小,各生境间差异显著。随光照强度降低,朝鲜淫羊藿的净光合速率、水分利用效率、气孔限制值、叶绿素a/b降低,而胞间CO2浓度、光能利用效率、叶绿素b和总叶绿素含量增加。各器官生物量积累随光照强度减弱而降低,各生境间差异极显著,其中,林缘地上生物量是林窗的1.6倍,林下的3.1倍。生物量分配方面,叶生物量比、叶面积比率、比叶面积随光照强度减弱而增加,根状茎生物量比和根冠比降低,林缘、林窗与林下差异极显著;林下、林缘、林窗朝鲜淫羊藿地上部分生物量比分别为0.733、0.659、0.664,林下朝鲜淫羊藿与林缘、林窗存在不同的光利用策略和生存策略。朝鲜淫羊藿作为极易遭受利用和生境破坏威胁的物种,宜在适度荫闭的林缘和疏林下开展野生抚育或仿生态栽培。     

不同光环境对桤木幼苗生长和光合特性的影响

通过设置3个光照强度(100%、56.2% 和12.5%),模拟森林幼苗生长的旷地(采伐迹地)、林窗和林下光环境,研究不同光照强度对外来种台湾桤木和乡土种四川桤木幼苗的生长、光合特性以及生物量积累与分配的影响.结果表明： 低光环境限制了两种桤木幼苗形态指标的增长,适当遮荫的林窗环境比旷地更有利于幼苗的生长.台湾桤木幼苗具有较高的比叶面积和相对生长速率,较大的单叶面积、叶长、叶宽、株高和基径,较少的叶片数和较低的叶面积比、叶柄长.低光环境下,台湾桤木幼苗的最大净光合速率、光饱和点和表观光量子效率较高,光补偿点和暗呼吸速率较低.随着光照强度的降低,台湾桤木幼苗具有更高的根生物量比和更低的叶生物量比;四川桤木幼苗则相反,加剧了动物取食和机械损伤的风险.     

不同光环境对桤木幼苗生长和光合特性的影响

通过设置3个光照强度(100％、56.2％和12.5％),模拟森林幼苗生长的旷地(采伐迹地)、林窗和林下光环境,研究不同光照强度对外来种台湾桤木和乡土种四川桤木幼苗的生长、光合特性以及生物量积累与分配的影响.结果表明:低光环境限制了两种桤木幼苗形态指标的增长,适当遮荫的林窗环境比旷地更有利于幼苗的生长.台湾桤木幼苗具有较高的比叶面积和相对生长速率,较大的单叶面积、叶长、叶宽、株高和基径,较少的叶片数和较低的叶面积比、叶柄长.低光环境下,台湾桤木幼苗的最大净光合速率、光饱和点和表观光量子效率较高,光补偿点和暗呼吸速率较低.随着光照强度的降低,台湾桤木幼苗具有更高的根生物量比和更低的叶生物量比;四川桤木幼苗则相反,加剧了动物取食和机械损伤的风险.     

不同光强对入侵种三裂叶豚草表型可塑性的影响

通过人工遮光,研究了不同光照强度下入侵植物三裂叶豚草的形态、生物量分配以及光合特性的表型可塑性.结果表明:与对照相比,遮光条件下,三裂叶豚草的株高、冠宽、单株叶面积、比叶面积和叶生物量比重显著增加,总生物量、单位叶面积生物量和根冠比减小;在全光照条件下,其冠宽和单株叶面积较小,根冠比较大,有利于高温强光下减少水分散失,表现出对不同光强较强的形态和生物量可塑性.遮光使三裂叶豚草叶片的日均净光合速率、蒸腾速率和气孔导度下降,胞间CO2浓度上升.正午光照最强时,低遮光处理植株的净光合速率、蒸腾速率和气孔导度最大.中遮光和高遮光条件下,其叶绿素含量显著增加,叶绿素a/b显著减小,有利于提高三裂叶豚草的光能利用效率,以适应弱光环境.     

麻栎和闽楠幼苗叶功能性状及生物量对光照和施肥的响应

光照和养分条件是影响植物生长的重要环境因子,不同生活型植物对环境异质性的响应机制不同。以落叶阔叶树种麻栎和常绿阔叶树种闽楠幼苗为研究对象,设置2个光照梯度（全光照和45%全光照）和4个施肥梯度（不施肥、氮磷供应比为5、15和45）共8种处理,研究光照和施肥及其交互作用对麻栎和闽楠生物量和叶形态、生理及化学性状的影响,并探讨了叶功能性状和生物量的关系。结果表明：（1）光照、施肥及其交互作用对光合气体交换参数（除水分利用效率外）、叶绿素荧光参数、叶形态指标（除比叶面积外）、单位质量叶氮含量和根冠比影响显著（P<0.05）。此外,光照和施肥对地上生物量和总生物量影响显著（P<0.05）。（2）全光照显著增加了麻栎和闽楠单株总叶面积和地上、地下生物量及总生物量（P<0.05）,而遮荫降低了非光化学猝灭系数、光合氮利用率和根冠比,增加了单位质量叶氮含量。（3）在全光照处理中,施肥显著增加了麻栎和闽楠水分利用效率（P<0.05）;在遮荫处理中,氮磷供应比45显著增加了麻栎和闽楠净光合速率和水分利用效率（P<0.05）。（4）麻栎和闽楠在全光照中倾向于资源获取策略,在遮荫中偏向于资源保守策略。在光照和施肥处理中,麻栎和闽楠单株总叶面积与地上生物量均显著正相关（P<0.05）。总之,单株总叶面积是预测麻栎和闽楠幼苗地上生物量变化的稳定指标,施肥有助于增加低光环境下麻栎和闽楠幼苗的生态适应能力。     

遮荫对3年生东北铁线莲生长特性及品质的影响

采用人工遮荫的方法,研究了遮荫处理(全光照CK、遮荫30%、50%、70%和90%)对东北铁线莲生长特性和品质的影响。结果表明:东北铁线莲株高随着遮荫梯度的增加而增高;30%遮荫处理茎粗最粗,根茎生物最高,与其它处理差异显著;而茎生物量、叶生物量则是30%遮荫处理与CK、70%和90%处理差异显著;东北铁线莲各器官生物量对光照强度的响应不同,表现为根茎生物量比、根冠比随着光照强度的减弱而降低,茎生物量比和叶生物量比则随着光照的减弱而增加,说明在遮荫的条件下,东北铁线莲优先将生物量分配给茎和叶,用来增加叶面积,以便于接受更多的光能,维持自身的生长发育。东北铁线莲光合色素含量随着遮荫梯度的增加而增加,90%遮荫下的东北铁线莲叶片光合色素含量最高,与其他处理差异显著;相反叶绿素a/b的值却最低;尤其是3年生东北铁线莲叶片中叶绿素 b含量随着光照强度的减弱而升高,能够使东北铁线莲充分吸收散射光,保持较强的光合能力。30%遮荫处理东北铁线莲净光合速率为10.86 μmol · m^-2 · s^-1,显著高于其它处理;随着光照强度的减弱东北铁线莲Gs先增加后降低,Ci则是先降低后增加,30%遮荫处理与其它处理间差异显著,但随着遮荫梯度的增加,东北铁线莲的净光合速率降低,光合作用合成产物总量减少,导致70%和90%遮荫条件下生物量较低;遮荫不利于东北铁线莲有效成分齐墩果酸的积累,不同处理,东北铁线莲齐墩果酸含量差异较大,以全光照和30%遮荫处理齐墩果酸含量较高,与其它处理间差异显著。因此,在生产实践上,对其进行适度遮荫,既保证东北铁线莲药材的质量,又提高药材产量;同时,若不以采收种子为目的,建议在8月份以后采收东北铁线莲药材。     

遮荫对闽楠幼苗光合及其叶片解剖特性的影响

以2年生闽楠幼苗为材料,探讨不同遮荫处理［0(CK,全光照) 和30%、50%、70%遮荫］对闽楠［Phoebe bournei (Hemsl.) Yang］幼苗光合作用及其叶片解剖结构的影响,为闽楠的合理栽培及人工林混交树种的配置提供理论依据。结果表明：(1)与全光照处理(CK)相比,闽楠幼苗叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(G_s)、最大净光合速率(P_nmax) 、光饱和点(LSP)、光补偿点(LCP)及暗呼吸速率(Rd)均随遮荫强度的增加而显著下降;在30%和50%遮荫下,幼苗能通过提高水分利用效率(WUE)和表观量子效率(AQY),增加叶绿素含量来充分利用弱光环境;70%遮荫处理则影响闽楠幼苗的正常生长。(2)遮荫降低了闽楠幼苗叶片的气孔密度,但单个气孔器的面积显著增加;遮荫处理下,闽楠叶片和主脉明显变薄;栅栏组织和海绵组织厚度降低,排列稀疏,栅海比减小,有利于提高对光能的利用效率。(3)闽楠幼苗根、茎、叶及总生物量均在全光照下最大,遮荫抑制了植株生物量的积累。研究发现,闽楠幼苗在遮荫处理下生物量积累和光合效率降低,但同时在叶片解剖结构和功能上表现出对遮荫的适应性,从而表现出一定的耐荫性。     

光强对杉木幼苗形态特征和叶片非结构性碳含量的影响

选取南方重要的造林树种杉木(Cunninghamia lanceolata(Lamb.)Hook)幼苗为研究对象,通过搭建遮荫棚设置5个光照强度(分别为自然光照的100%、60%、40%、15%和5%),研究了幼苗在不同光照强度下的生长形态、生物量积累及分配、叶片的非结构性碳含量(NSC)特征。结果显示:(1)叶长、叶宽和叶面积在40%光照强度下最大,而比叶面积和叶片相对含水量随着光照强度的降低呈递增趋势;(2)随着光照强度的降低,杉木幼苗各器官生物量下降,根生物量比和根冠比降低,茎和叶生物量比增加;(3)杉木幼苗在60%光照强度下叶片非结构性碳含量最高,5%光照强度下含量最低;(4)杉木幼苗比叶面积与叶生物量以及与非结构性碳含量之间存在极显著的负相关关系(P0.01),叶生物量与非结构性碳含量之间存在极显著的正相关关系(P0.01)。杉木幼苗能够通过形态学上的可塑性来适应不同的光强环境,提高光竞争能力和生存适合度,但在5%光照强度下,由于较难维持碳收支平衡而不利于其生长和存活。     

披针叶茴香对变化光环境的表型可塑性

植物对变化光环境的表型可塑性大小影响其在林下生境中分布、生长和更新。为探讨披针叶茴香在不同光环境下的整体表型可塑性及其适应机制,采用遮荫试验模拟5种光照条件(100%、52%、33%、15%和6%相对光照强度),研究了不同光环境下披针叶茴香叶片形态、生理、解剖结构、根系形态以及生物量分配等的变化。结果表明:叶生物量在5种光照处理之间差异不显著,但叶面积和比叶面积均随光照强度减弱显著增加。遮荫处理增加了叶绿素a、叶绿素b和类胡萝卜素的含量,但叶绿素a/b比值随光照强度减弱而降低。遮荫降低了非结构性碳水化合物(淀粉和可溶性糖)和可溶性蛋白的含量,增加了叶片氮和磷含量,对叶片氮/磷比影响较小。在52%和33%相对光照处理下,叶片中硝酸盐含量最低,而在100%和6%相对光照处理下硝酸盐积累较多。根生物量、细根和粗根的长度、表面积以及比根长和比根表面积在5种光照处理之间均没有显著差异,根系氮含量在低光环境(15%和6%相对光照处理)中显著降低。随光照强度减弱,披针叶茴香采取保守生存策略,并没有增加叶生物量的分配,而是分配较多的生物量给枝条和树干,储存能量。综合来看,披针叶茴香具有较宽的光生态幅,在6%—100%光照强度下均能正常生长,遮荫有利于披针叶茴香地上和总生物量的积累,52%的相对光照条件下生长最佳。变化光环境下根系性状和整体结构的可塑性相对较低,叶片生理性状的可塑性在披针叶茴香适应光环境变化过程中发挥了主要作用。     

光照对鄂东南2种落叶阔叶树种幼苗生长、光合特性和生物量分配的影响

通过设置4个光照梯度(25%、12%、6%和3%自然光)模拟鄂东南低山丘陵地区落叶阔叶林林下的光环境,研究了2种耐荫性不同的树种幼苗--麻栎(Quercus acutissima)和化香(Platycarya strobilacea)不同光强下的存活率、光合特性、生长和生物量分配,探讨了低光环境中耐荫性不同的树种幼苗维持自身碳平衡的机制和权衡"存活-生长"选择的生活史策略。结果表明:(1)低光下的2个树种幼苗的生长、光合特性和生物量分配具有显著性差异。(2)各个光照梯度下麻栎幼苗都生长良好,存活率保持在35%以上,而化香幼苗遭遇高的死亡率,80d后3%和6%自然光下的幼苗全部死亡;低光环境中麻栎幼苗比化香幼苗具有更大的表观光量子(AQY)和最大净光合效率(Pmax),更低的光补偿点(LCP)和暗呼吸效率(Rd),即耐荫性较强的麻栎幼苗比耐荫性较弱的化香幼苗具有更高的低光碳同化率和碳捕获能力。(3)2个树种幼苗的成活率与RGR呈负相关关系,各个光照梯度下耐荫性较弱的化香幼苗的相对生长率(RGR)显著高于耐荫性较强的麻栎幼苗,而两个树种幼苗的净同化率(NAR)无明显差异。相对于麻栎幼苗较高的根生物量比(RMR),化香幼苗将更多的生物量分配给叶部,因而具有较高的叶生物量比(LMR)、叶面积比(LAR)和比叶面积(SLA)。不同耐荫性的幼苗生长及生物量分配方式的差异是植物"存活-生长"权衡后的结果,耐荫性弱的化香幼苗具有较高的生长潜力和较弱的自我保护能力,而耐荫性强的麻栎幼苗具有更高的低光碳储量,能够维持更好的低光碳平衡,具有竞争优势。     

苗圃氮加载及水分胁迫对栓皮栎苗木叶片光合氮分配及生物量积累的影响

苗圃科学施氮(N)作为提高苗木N贮存水平与质量的核心手段,能否提高干旱立地苗木造林效果仍存在争议;N贮存水平与干旱如何协同作用影响叶片光合N分配及苗木生物量积累尚不明确。阐明上述问题,能够为干旱立地下的森林植被恢复以及造林苗木科学精准施N提供科学依据。选择栓皮栎(Quercus variabilis Blume)为研究对象,对一年生苗木设置2个苗圃木质化期N加载水平(0、24 mg N/株),翌年春苗木移栽后设置2个灌溉水平(85%、40%田间持水量),取样测定苗木生物量、叶片N、叶绿素与脯氨酸水平、以及气体交换参数,计算光合N分配及光合N利用效率(PNUE)。结果表明,叶片发育完成后,干旱抑制N向光合系统分配,但N加载处理提高了干旱下的光合N含量,从而在一定程度上抵消干旱对生物量积累的抑制;无N加载苗木则向光合系统投入更少的N,而提高脯氨酸水平,生物量积累受抑制更为显著。无N加载苗木在遭受干旱后将N向羧化组分分配,而N加载苗木遭遇干旱后则显著抑制叶片将N向羧化系统以及电子传递系统分配,捕光组分N的分配则不受植物体内N贮存或外部水分状况的影响,栓皮栎苗木通过调整不同功能组分光合N含量和...     

Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size

An evergreen oak species, Cyclobalanopsis multinervis, and a deciduous oak species, Quercus aliena var. acuteserrata were grown from acorns under two light levels (full sunlight and shade at about 18 % of full sunlight, simulating the light intensities in forest clearings and gaps, respectively) for one growing season. Three hypotheses were tested: (i) the deciduous species grows faster than the evergreen species in forest gaps and clearings; (ii) the deciduous species responds more strongly in terms of growth and morphology to variation in light climate than the evergreen species; and (iii) seedling size is positively correlated to acorn size. The results showed: (i) at both light levels, the deciduous seedlings gained significantly more growth in biomass and height than the evergreen seedlings; (ii) both species produced significantly more biomass in full sunlight than in shade, without showing any significant difference in height between treatments. Increase in light intensity improved the growth of the deciduous seedlings more strongly; (iii) at a similar age, the deciduous seedlings showed a greater response in leaf morphology and biomass allocation to variation in light levels, but when compared at a similar size, biomass allocation patterns did not differ significantly between species; (iv) bigger acorns tended to produce larger seedlings, larger leaf sizes and more leaf area, between and within species. These differences demonstrate that the deciduous species is gap-dependent and has the advantage over the evergreen species in forest gaps and clearings.     

不同光强对棱角山矾幼苗生长及光合特性的影响

通过比较不同遮阴处理下棱角山矾幼苗生长及光合特性的差异,以探讨不同光强对棱角山矾生长及光合作用的影响及其适应机制。结果表明：棱角山矾幼苗的生物量分配对光强的反应敏感,在强光处理下,比叶面积减小,总叶面积变小;弱光处理下通过增大叶生物量比、叶面积比,支持结构生物量比,提高捕光能力。但L100、L30、L10处理下总生物量的积累显著小于L50处理,L30、L50、L100光强处理下总叶面积显著高于L10光强处理。并且10%和100%光强下棱角山矾叶片的最大净光合速率（P_max）、光饱和点（LSP）、表观量子效率（AQY）均显著低于30%和50%光强处理;而光补偿点（LCP）则随光强的增大而升高;总叶绿素和类胡萝卜素含量在10%~100%随着光强增大而显著降低（P<0.05）。说明棱角山矾幼苗在不同光环境下具有一定适应性,但过强或过弱的光环境均会对棱角山矾的生长产生负面影响,其中50%光强是棱角山矾幼苗生长的最佳光强;棱角山矾为中性树种,苗木适宜生长在荫蔽的环境下。     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Biomass allocation, shade tolerance and seedling survival of the invasive species

Berberis darwinii (Berberidaceae) is a serious environmental weed in New Zealand, capable of invading a range of different light environments from grazed pasture to intact forest. According to optimal partitioning models, some plants optimise growth under different environmental conditions by shifting biomass allocation among tissue types (e.g. roots, shoots) to maximise the capture of limiting resources (e.g. water, light). We examined patterns of growth, biomass allocation, and seedling survival in Berberis darwinii to determine whether any of these factors might be contributing to invasion success. Growth and biomass allocation parameters were measured on seedlings grown for 7 months in five natural light environments in the field. Survival was high in the sunniest sites, and low in the shadiest sites. Seedlings grown in full sun were an order of magnitude taller and heavier, had five times as many leaves, and proportionally more biomass allocated to leaves than seedlings grown in other light environments. In the shade, leaves were bigger and thinner, and leaf area as a proportion of total plant biomass increased, but the proportion of above- to below-ground biomass was similar across all light and soil moisture environments. In summary, although leaf traits were plastic, patterns of biomass allocation did not vary according to optimal partitioning models, and were not correlated with patterns of seedling survival. Implications for the management of this invasive species are discussed.     

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.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.     

Why is it better to produce coffee seedlings in full sunlight than in the shade? A morphophysiological approach

The coffee plant is native to shaded environments and its seedlings are often produced in shaded nurseries. However, some nursery managers, in an effort to improve the acclimation of seedlings to field conditions after transplantation, produce seedlings in full sun exposure. In this study, the morphological and physiological parameters of arabica coffee (Coffea arabica) seedlings produced in full sun (T1) and in shade (T2) were examined. The biomass accumulation and relative growth rate of T1 and T2 seedlings were similar. The T1 seedlings had less biomass allocation to shoots, a lower leaf mass ratio and a lower leaf area ratio; however, they had a greater net assimilation rate (rate of increase in plant mass per unit leaf area), which was associated with a greater net photosynthetic rate. There were no alterations in the concentrations of total chlorophylls or in the chlorophyll a/b ratio when comparing T1 and T2 seedlings. No indications of photoinhibition or photooxidative damage were observed in the T1 plants, which were shown to have a more robust antioxidant system than the T2 plants. Seedlings transferred from shade to full sun (T3) were not capable of utilising the incident extra light to fix CO₂. These seedlings showed a remarkable nocturnal retention of zeaxanthin and a significantly increased deepoxidation state of the xanthophyll cycle, even at predawn, but the activity of antioxidant enzymes was lower than in the T1 and T2 plants. Despite the acclimation capacity of T3 seedlings to the new light environment, they exhibited chronic photoinhibition and considerable photooxidative damage throughout the seven days following the transfer to full sun exposure. We further discuss the practical implications of producing coffee seedlings in full sunlight and under shade.