西双版纳热带雨林中两种生态型蕨类植物的光合特性比较研究

通过比较分布于西双版纳热带雨林林下生境中的附生鸟巢蕨(Neottopteris nidus)和地生网脉铁角蕨(Asplenium finlaysonianum)的光合特征和光合诱导特性, 来研究不同生态型蕨类植物的光斑利用策略。研究结果表明, 2种蕨类植物的最大净光合速率、暗呼吸速率、表观量子效率、光饱和点和光补偿点没有显著差异, 但网脉铁角蕨的最大气孔导度远远高于鸟巢蕨, 表明后者具有更强的光合水分利用效率。在暗处理3小时接着光照(光强为20 mmol .m-2.s-1)30分钟后, 网脉铁角蕨的初始气孔导度显著高于鸟巢蕨。连续照射饱和强光后, 网脉铁角蕨达到最大净光合速率50%(T50%)和90%的时间(T90%)比鸟巢蕨短: 网脉铁角蕨和鸟巢蕨的T50%分别为0.57和5.31分钟, T90%分别为5.85和26.33分钟。诱导过程中, 气孔导度对强光的响应明显滞后于净光合速率。鸟巢蕨达到最大气孔导度的时间明显比网脉铁角蕨慢, 但在光合诱导消失过程中2种蕨类植物的 光合诱导维持能力却没有显著差异。上述结果表明, 与大多数地生林下植物(如网脉铁角蕨)相比, 附生鸟巢蕨的水分保护比碳获得更重要, 但却限制了附生蕨对光斑的利用。     

6种热带雨林木本植物幼苗光合诱导的研究

 在晴天上午适宜条件下,测定了生长在模拟林下光环境中的6种热带雨林木本植物幼苗的光合特性和光合诱导特征。6种植物分别为先锋树种大穗野桐(Mallotus macrostachys),冠层树种绒毛番龙眼(Pometia tomentosa)、玉蕊(Barringtonia pendala)、望天树(Shorea chinensis)、滇南插柚紫(Linociera insignis)和林下灌木睫毛粗叶木(Lasianthus hookeri)。研究结果表明：暗处理3 h的叶片经连续饱和强光照射后,6种植物的净光合速率呈s形到双曲线形。6种植物达到90%最大净光合速率的时间为4.4～12.5 min,这与所报道的其它热带雨林中一些阴生植物的诱导速率相近。大穗野桐和睫毛粗叶木的诱导速率最快,达到50%和90%最大净光合速率的时间为其它4种冠层植物幼苗的1/2至1/3。诱导过程中,最大气孔导度对强光的响应明显滞后于净光合速率。充分诱导的叶片在黑暗中20 min后,6种植物的诱导状态都较高。其中,大穗野桐的诱导状态消失相对较快,这可能与其气孔导度和羧化能力的快速降低有关。玉蕊诱导状态的消失主要与生化限制有关,因为此时它的气孔导度仍维持相对较高的值。而睫毛粗叶木较高的气孔导度和羧化能力的维持导致了很高的诱导状态。林下植物这种对强光的快速反应和黑暗中高的维持状态对有效利用光斑具有重要的意义,这与其一生中在林下生长和更新的特点是一致的。     

北京松山油松当年生与往年生针叶光合生理特性

为探讨北京松山自然保护区油松固碳机制, 对松山不同叶龄油松针叶的光合、蒸腾特性及其与环境因子的关系进行探究。结果显示：(1)油松当年生叶的最大净光合速率、暗呼吸速率及光补偿点均高于往年生叶, 但表观量子效率却低于往年生叶; (2)油松当年生和往年生叶的净光合速率和蒸腾速率日变化均呈“双峰”曲线; 当年生叶的净光合速率日均值显著高于往年生(P<0.05); (3)气孔导度和光合有效辐射是影响油松当年生和往年生叶净光合速率的共同生态因子; 同时气孔导度和光合有效辐射共同影响不同叶龄针叶的蒸腾速率。结果表明当年生叶光合潜力较大, 而往年生叶利用弱光能力较强; 当年生叶净光合速率降低主要为非气孔因素, 而往年生叶则先后由气孔因素和非气孔素引起; 油松不同叶龄光合特性差异体现了油松不 同的生理调节和环境适应策略。     

虎杖光合生理生态特性日变化研究

用Li-6400便携式光合测定系统对5个虎杖材料的光合生理特性日变化及其与气象因子关系进行了研究。结果表明：（1）虎杖的净光合速率日变化呈‘单峰’型曲线,日最大净光合速率（15．0μmol·m^-2·s^-1）值出现在9：00左右;（2）叶片水压亏缺、气孔导度和蒸腾速率的峰值在同一时间出现（13：00）,胞间CO2浓度不随气孔导度的降低而减小,控制虎杖光合速率因子为非气孔限制;（3）供试5个材料的净光合速率日变化趋势基本一致,并以地栽组培苗的净光合速率最高,而‘贵州凯里’的最低。光合有效辐射对光合特征参数的变化影响最大,且对净光合速率起决定性作用（r-0．534^＊＊）,环境因子主要通过对蒸腾速率、叶片水压亏缺和叶面温度的作用来影响虎杖叶片净光合速率。     

沈阳城市森林主要绿化树种——丁香的光合特性研究

应用LI-6400红外气体分析仪对城市森林典型树种丁香不同季节光合作用的日变化规律进行研究,采用相关分析和多元逐步回归统计分析方法,得出净光合速率与光合有效辐射、气温、大气湿度、气孔导度等生理生态因子间的相关关系．结果表明,丁香春季和夏季光合速率呈双峰曲线,秋季呈单峰曲线．春季影响光合速率的主要因子为光合有效辐射和气孔导度;夏季影响光合速率的主要因子为气孔导度和光合有效辐射;秋季为气孔导度和胞间CO2浓度,建立了相应的影响因子回归方程。     

二倍体小麦种间气孔与光合特征差异

选取5个二倍体小麦种为实验材料,利用数码图像显微镜处理系统、气体交换参数测定系统和叶绿素荧光动力学参数测定系统,研究了二倍体小麦种间气孔特征与光合特性的差异.结果表明,二倍体小麦种间气孔和光合特征存在较大的差异:其中S基因型(Ae.Sect.Sitopsis)的气孔长度、周长和面积最大,而其宽度、密度和气孔指数却是最小的;D基因型(Ae.Tauschii)的气孔宽度、密度和气孔指数最大,而其长度、周长和面积却是最小的;D基因型(Ae.Tauschii)的净光合速率最大,而S基因型(Ae.Sect.Sitopsis)的净光合速率最小.气孔周长与气孔面积呈显著正相关关系,而气孔密度与气孔面积呈显著负相关关系.二倍体小麦的净光合速率与气孔导度呈显著正相关关系,同时气孔限制值的变化趋势与细胞间隙CO2浓度的变化趋势相反,说明二倍体小麦的光合能力主要受气孔限制.在几个种中,乌拉尔图小麦(T.urartu)具有较高的净光合速率和气孔导度,同时细胞间隙CO2浓度最低,气孔限制值最大,在光合能力和耐光抑制方面有较明显的优势;而野生一粒小麦(T.boeoticum)则在光合能力和耐光抑制方面不存在优势.同时,乌拉尔图小麦(T.urartu)的叶绿素含量最高,而野生一粒小麦(T.boeoticum)最低,叶绿素含量可能也是造成二倍体种间光合能力差异的原因之一.     

沈阳城市森林主要绿化树种——丁香的光合特性研究

应用LI-6400红外气体分析仪对城市森林典型树种丁香不同季节光合作用的日变化规律进行研究,采用相关分析和多元逐步回归统计分析方法,得出净光合速率与光合有效辐射、气温、大气湿度、气孔导度等生理生态因子间的相关关系.结果表明,丁香春季和夏季光合速率呈双峰曲线,秋季呈单峰曲线.春季影响光合速率的主要因子为光合有效辐射和气孔导度;夏季影响光合速率的主要因子为气孔导度和光合有效辐射;秋季为气孔导度和胞间CO₂浓度.建立了相应的影响因子回归方程.     

银杏叶片光合作用对强光的响应

银杏叶片遭受光量子通量密度（PFD）为1200μmolm-2s-1的强光胁迫后,净光合速率（Pn）、气孔导度（Gs）、细胞间隙CO2浓度（Ci）、PSⅡ光化学效率（Fv／Fm）和表现量子效率（AQY）都下降,而叶片在505nm处的光吸收（A505）、初始荧光水平（Fo）和荧光的非光化学猝灭（qN）上升。在去除强光胁迫数小时之后,这些参数都不能完全恢复。这就表明,虽然强光能引起严重的光抑制,可能涉及依赖叶黄素循环的热耗散和一部分PSⅡ反应中心的失活及破坏,但是导致光合速率降低的主要因素仍然是气孔导度的降低。     

沙坡头主要建群植物油蒿和柠条的气体交换特征研究

在自然条件下对沙坡头人工固沙区主要建群植物油蒿和柠条的气体交换特征进行了比较研究。结果表明,油蒿和柠条都具有典型的双峰型光合速率日进程曲线,它们的气孔导度日变化趋势也与净光合速率日变化相一致;在一日内油蒿的气孔导度、净光合速率和蒸腾速率始终大于柠条。油蒿和柠条在８：００￣１０：００光合速率下降的主要决定因素是气孔导度,而在午间（１０：００￣１４：００）和午后（１６：００￣１９：００）,光合速率的上     

遮荫对东北地区四种可食用蕨类植物生长和光合特征的影响

以东北4种蕨类植物——荚果蕨(Matteuccia struthiopteris(L.)Todaro)、猴腿蹄盖蕨(Athyrium multidentatum(Dol1.)Ching)、分株紫萁(Osmunda cinnamomea(L.)var.asiatica Fernald)和蕨(Pteridium aquilinum (L.) Kuhn var.latiusculum (Desy.) Underw.ex Heller)为研究材料,分析在35%、13%和8%全光照处理下4种蕨类植物的生长和光合特征.结果发现:随着遮荫程度的增加,4种蕨类植物的株高减小,比叶面积增加,叶片含水量增加.4种蕨类植物的净光合速率、气孔导度、蒸腾速率随着遮荫程度的增加而降低,胞间CO2浓度呈上升趋势.在3种遮荫处理下4种蕨类植物的PSⅡ最大光化学效率在0.7~0.8;非光化学猝灭系数随着遮荫程度的增加而降低.4种蕨类植物的叶绿素含量,尤其是叶绿素b含量随着遮荫程度的增加而上升,但在8%全光照下蕨的叶绿素含量呈下降趋势.相比其他蕨类植物而言,蕨具有较高的比叶面积、净光合速率和量子产率,相对较低的非光化学猝灭系数和叶绿素a./b.结果表明,4种蕨类植物最适合在35%全光照下生长,虽然对较低光照条件具有一定的适应性,但生长受到一定的抑制;4种蕨类植物中,蕨具有较强的耐荫性.本研究为东北地区4种可食用蕨类植物的栽培管理及利用提供科学依据.     

Leaf Photosynthetic Rate of Tropical Ferns Is Evolutionarily Linked to Water Transport Capacity

Ferns usually have relatively lower photosynthetic potential than angiosperms. However, it is unclear whether low photosynthetic potential of ferns is linked to leaf water supply. We hypothesized that there is an evolutionary association of leaf water transport capacity with photosynthesis and stomatal density in ferns. In the present study, a series of functional traits relating to leaf anatomy, hydraulics and physiology were assessed in 19 terrestrial and 11 epiphytic ferns in a common garden, and analyzed by a comparative phylogenetics method. Compared with epiphytic ferns, terrestrial ferns had higher vein density (D_vein), stomatal density (SD), stomatal conductance (g_s), and photosynthetic capacity (A_max), but lower values for lower epidermal thickness (LET) and leaf thickness (LT). Across species, all traits varied significantly, but only stomatal length (SL) showed strong phylogenetic conservatism. A_max was positively correlated with D_vein and g_s with and without phylogenetic corrections. SD correlated positively with A_max, D_vein and g_s, with the correlation between SD and D_vein being significant after phylogenetic correction. Leaf water content showed significant correlations with LET, LT, and mesophyll thickness. Our results provide evidence that A_max of the studied ferns is linked to leaf water transport capacity, and there was an evolutionary association between water supply and demand in ferns. These findings add new insights into the evolutionary correlations among traits involving carbon and water economy in ferns.     

Photosynthetic response of four fern species from different habitats to drought stress: relationship between morpho-anatomical and physiological traits

Ferns flourish in many habitats, from epiphytic to terrestrial and from sunny to shady, and such varied conditions require contrasting photosynthetic strategies to cope with drought. Four species of temperate ferns from different habitats were subjected to drought by withholding irrigation in order to investigate their photosynthetic responses. Lepisorus thunbergianus (epiphytic) had low stomatal density and showed high water-use efficiency (WUE) retaining photosynthetic activity with low relative frond water content under drought stress, which suggested their high adaptation to drought. On the other hand, low WUE with low light-saturated photosynthetic rate in Adiantum pedatum (terrestrial, shady environment) was associated with much lower photosynthesis than in the other species under drought stress, suggesting lower adaptation to drought-prone habitats. Morphological stomatal traits such as stomatal density and photosynthetic response to drought in ferns involved species-specific adaptation to survive and grow in their natural habitats with different levels of drought.     

Seedling Mortality in Hawaiian Rain Forest: The Role of Small‐Scale Physical Disturbance1

Most montane rain forests on the island of Hawaii consist of a closed canopy formed by Cibotium spp. tree ferns beneath an open canopy of emergent Metrosideros polymorpha trees. We used artificial seedlings to assess the extent to which physical disturbance caused by the senescing fronds of tree ferns and the activities of feral pigs might limit tree regeneration. Artificial seedlings were established terrestrially (N= 300) or epiphytically (N = 300) on tree fern stems. Half of the seedlings on each substrate were in an exclosure lacking feral pigs and half were in forest with pigs present. After one year, the percentage of seedlings damaged was significantly greater among terrestrial seedlings (25.7%) than epiphytic seedlings (11.3%). Significantly more terrestrial seedlings were damaged in the presence of pigs (31.3%) than in the absence of pigs (20.0%). Senescing fronds of tree ferns were responsible for 60.3 percent of the damaged seedlings. Physical disturbance is potentially a major cause of seedling mortality and may reduce the expected half‐life of a seedling cohort to less than two years.     

Amazonian Dark Earth Shapes the Understory Plant Community in a Bolivian Forest

Amazonian Dark Earths (ADE) are the result of human modification of the Amazonian landscape since pre‐Columbian times. ADE are characterized by increased soil fertility compared to natural soils. In the Amazonian forest, soil fertility influences understory herb and fern species composition. However, little research has been done to evaluate the effect of ADE on the composition of the understory community. We evaluated the effects of ADE and soil in 36 plots (150 m × 4 m) established in a Bolivian moist forest (La Chonta). For each plot, we determined soil nutrients, and the composition, richness, and abundance of terrestrial fern, angiosperm herb, and understory palm species. We found that the presence of ADE created a gradient in soil nutrients and pH that affected the understory species composition especially of ferns and palms. Additionally, the higher nutrient concentration and more neutral soil pH on ADE soils caused a decrease of ferns species richness. We therefore conclude that the current composition of the understory community in this particular Bolivian forest is a reflection of past human modifications of the soil.     

The influence of life form on carbon and nitrogen relationships in tropical rainforest ferns

Tropical ferns are characterized by a high diversity of plant life forms, yet there have been few large-scale studies on the functional ecology of these different forms. We examined epiphytic, hemiepiphytic, and terrestrial ferns, and asked whether there are differences in the mineral nutrition and water relations across different growth forms of a diverse assemblage of species. We measured specific leaf area, leaf nitrogen concentrations, and natural abundance of the stable isotopes δ¹⁵N and δ¹³C of 48 fern species from 36 genera across a wide range of habitats at La Selva Biological Station in Costa Rica. We found that epiphytes were significantly different in all measured variables from hemiepiphytic and terrestrial species, and that terrestrial and soil-rooted hemiepiphytes were indistinguishable in all variables excluding SLW. A multivariate analysis revealed that aspects of N nutrition were the most reliable at separating epiphytic species from other life forms. Our study demonstrates that the natural abundance of both C and N as well as N relations and leaf morphology are useful when segregating different plant life forms, and that the N cycle of epiphytic and terrestrial habitats function independently from each other.     

Genetic diversity and breeding system in a group of neotropical epiphytic ferns (Pleopeltis; Polypodiaceae)

Epiphytes are ecologically important components of tropical forests worldwide and yet they have been underrepresented in studies of reproductive biology. Given the presumed ephemeral nature of their substrates, and the importance of dispersal and colonization, epiphytes might be expected to undergo substantial inbreeding to ensure reproductive success, as in weedy terrestrial plants. While there is some evidence for inbreeding in epiphytic angiosperms, the only previous studies of fern epiphytes indicate that they are predominantly outcrossing. The present study reports on the genetic diversity and breeding system of six members of the Neotropical epiphytic fern genus Pleopeltis (Polypodiaceae). A survey of isozyme variability using starch gel electrophoresis revealed high population levels of polymorphism (P = 0.62), allelic diversity (A = 2.3), and individual heterozygosity (Ho = 0.181), but little differentiation among conspecific populations (I 3 0.98; Gst = 0.048), and high interpopulational gene flow rates (Nm > 1). In addition, there was no indication of homozygote excess within populations that might indicate a history of selfing in these ferns: populations generally conformed to Hardy-Weinberg expected genotype frequencies, and both Wright's inbreeding coefficient (Fis) and Holsinger's intragametophytic selfing rates approached zero. Possible mechanisms limiting inbreeding in these ferns include antheridiogen activity and high levels of genetic load that would lead to inbreeding depression upon selfing.     

鼎湖山季风常绿阔叶林不同生境蕨类植物的叶片功能性状研究

为了解不同生境中蕨类植物的叶片功能,对鼎湖山季风常绿阔叶林中的6种优势蕨类植物的比叶面积、光合速率、元素含量、构建成本和水分利用效率等叶片性状进行了测定,研究了它们对环境的适应性。研究结果表明,两种附生蕨类(江南星蕨和崖姜蕨)的叶片具有较低的比叶面积(SLA)和光合速率(Aarea),较高的构建成本(CCarea)和碳同位素比率(?13C)。而两种林下土生蕨类(扇叶铁线蕨、桫椤)具有较高的Aarea和较低的?13C。生长在岩石基质上的中华里白具有最高的养分利用效率和较高的?13C。生长在溪流边的薄叶卷柏具有最高的SLA,最低的CCarea和?13C。这些揭示了蕨类植物叶片性状与生境之间具有紧密的相关性。由于附生蕨类的叶片具有高的能量投资和低的光合速率,对环境变化的适应能力弱,附生蕨类可能面临较大的生存压力。     

Photosynthetic responses to dynamic light under field conditions in six tropical rainforest shrubs occuring along a light gradient

 We examined in the field the photosynthetic utilization of fluctuating light by six neotropical rainforest shrubs of the family Rubiaceae. They were growing in three different light environments: forest understory, small gaps, and clearings. Gas exchange techniques were used to analyse photosynthetic induction response, induction maintenance during low-light periods, and lightfleck (simulated sunfleck) use efficiency (LUE). Total daily photon flux density (PFD) reaching the plants during the wet season was 37 times higher in clearings than in the understory, with small gaps exhibiting intermediate values. Sunflecks were more frequent, but shorter and of lower intensity in the understory than in clearings. However, sunflecks contributed one-third of the daily PFD in the understory. Maximum rates of net photosynthesis, carboxylation capacity, electron transport, and maximum stomatal conductance were lower in understory species than in species growing in small gaps or clearings, while the reverse was true for the curvature factor of the light response of photosynthesis. No significant differences were found in the apparent quantum yield. The rise of net photosynthesis during induction after transfer from low to high light varied from a hyperbolic shape to a sigmoidal increase. Rates of photosynthetic induction exhibited a negative exponential relationship with stomatal conductance in the shade prior to the increase in PFD. Leaves of understory species showed the most rapid induction and remained induced longer once transferred to the shade than did leaves of medium- or high-light species. LUE decreased rapidly with increasing lightfleck duration and was affected by the induction state of the leaf. Fully induced leaves exhibited LUEs up to 300% for 1-s lightflecks, while LUE was below 100% for 1–80 s lightflecks in uninduced leaves. Both induced and uninduced leaves of understory species exhibited higher LUE than those of species growing in small gaps or clearings. However, most differences disappeared for lightflecks 10 s long or longer. Thus, understory species, which grew in a highly dynamic light environment, had better capacities for utilization of rapidly fluctuating light than species from habitats with higher light availability. Received: 4 January 1997 / Accepted: 28 April 1997