阔叶红松林3种典型蕨类叶功能性状的垂直变异
Vertical variations in leaf functional traits of three typical ferns in mixed broadleaved- Korean pine forest
通讯作者: *刘志理(liuzl2093@126.com)
编委: 吕晓涛
责任编辑: 乔鲜果
收稿日期: 2022-10-21 接受日期: 2023-03-27
基金资助: |
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Corresponding authors: *LIU Zhi-Li(liuzl2093@126.com)
Received: 2022-10-21 Accepted: 2023-03-27
Fund supported: |
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研究蕨类在不同垂直高度下叶功能性状及其相关关系的差异, 可为揭示大型蕨类叶片的资源利用策略提供科学依据。以阔叶红松(Pinus koraiensis)林内的粗茎鳞毛蕨(Dryopteris crassirhizoma)、东北蹄盖蕨(Athyrium brevifrons)和荚果蕨(Matteuccia struthiopteris) 3种蕨类为研究对象, 将个体按叶片垂直高度分为上、中、下3层, 分别测定其比叶面积、叶干物质含量、净光合速率、瞬时水分利用效率、叶氮含量和叶磷含量, 并测定每株个体的光环境和土壤因子, 重点揭示叶功能性状的垂直变异规律及相关关系。结果表明: (1) 3种蕨类叶干物质含量随叶片垂直高度增加均呈递增趋势, 比叶面积无显著差异; 东北蹄盖蕨和荚果蕨净光合速率随叶片垂直高度增加呈递增趋势; 荚果蕨瞬时水分利用效率随叶片垂直高度增加先增后减, 叶氮含量逐渐减少; 粗茎鳞毛蕨叶磷含量随叶片垂直高度增加表现为先增后减。(2)叶氮含量与比叶面积, 瞬时水分利用效率与净光合速率显著正相关; 叶氮含量与叶干物质含量, 叶干物质含量与比叶面积则显著负相关; 上述叶功能性状间的相关关系在不同垂直高度间均无显著差异。(3)土壤有效磷含量和土壤pH是不同垂直高度叶功能性状变异的主要影响因子, 土壤有效磷含量对叶功能性状变异的解释度最高。该研究表明, 阔叶红松林内大型蕨类的叶功能性状存在一定的垂直差异, 然而个体内部性状间的变化速率基本恒定, 光环境和土壤因子对不同垂直高度叶功能性状变异的影响程度不同。
关键词:
Aims Understanding the differences in leaf functional traits and their correlations in ferns at different vertical heights can provide a scientific basis for revealing the resource utilization strategies of large fern fronds.
Methods Individuals of three fern species in a mixed broadleaved-Korean pine (Pinus koraiensis) forest, i.e., Dryopteris crassirhizoma, Athyrium brevifrons and Matteuccia struthiopteris were divided into upper, middle, and lower layers according to the vertical height of leaves. We measured specific leaf area, leaf dry matter content, net photosynthetic rate, instantaneous water use efficiency, leaf nitrogen content and leaf phosphorus content, as well as the light environment and soil factors of each individual plant to reveal the vertical variation patterns and correlations of leaf functional traits.
Important findings (1) Leaf dry matter content of the three fern species increased with the vertical height of the fronds, but specific leaf area showed no variation. The net photosynthetic rate of A. brevifrons and M. struthiopteris showed an increasing trend with the increases of vertical height of fronds, the instantaneous water use efficiency of M. struthiopteris increased and then decreased with the vertical height of the fronds, and leaf nitrogen content gradually decreased; leaf phosphorus content of D. crassirhizoma showed an increase and then decreased trend. (2) There were positive correlations between leaf nitrogen content and specific leaf area, and also between instantaneous water use efficiency and net photosynthetic rate. There were negative correlations between leaf nitrogen content and leaf dry matter content, and between leaf dry matter content and specific leaf area. The correlations among those leaf functional traits did not differ significantly among different vertical heights. (3) Soil available phosphorus content and soil pH were the main factors affecting the variation of leaf functional traits at different vertical heights, with soil available phosphorus content having the highest explanatory degree to the variation of leaf functional traits. Our results indicated that there were vertical differences in leaf functional traits of large ferns in the mixed broadleaved-Korean pine forest, but the rate of change among individual characters was basically constant, the effects of light environment and soil factors on the variation of leaf functional traits differed among vertical heights. This study provided reference for further research on the mechanism of leaf functional traits variation in different vertical heights of ferns in understory.
Keywords:
引用本文
赵孟娟, 金光泽, 刘志理.
ZHAO Meng-Juan, JIN Guang-Ze, LIU Zhi-Li.
森林植物叶片的分布、大小和方向决定了冠层内光利用方式(Yu & Sun, 2013; Cheng et al., 2022), 同时, 冠层差异意味着不同垂直高度叶片光拦截和资源利用能力的变化(Osada & Hiura, 2017), 冠层开放度扩大, 林内的光照总辐射增多, 叶片截获光资源并充分利用叶绿素可吸收波段的光进行光合作用(李晖等, 2021), 表现为植物叶功能性状的垂直变异。比叶面积反映植物对光、温、水等资源的利用效率, 与叶氮含量密切相关(Reich et al., 1997; Roche et al., 2004; Wright et al., 2004); 干物质含量为叶片干质量与饱和鲜质量的比值, 一定程度反映了叶片对养分的获取和储存能力(Wilson et al., 1999); 净光合速率反映植物叶片光强利用率、耐阴性及光合作用效率, 是各尺度下植物生理生态学研究的重要指标(Pierangelini et al., 2014; Han et al., 2022); 水分利用效率表征植物在消耗等量水分时固定CO2的能力, 是衡量植物适应环境胁迫的关键指标(赵自国等, 2019); 氮是叶绿素的重要组成部分, 与植物光合作用密切相关(Reich et al., 1997); 叶氮、磷含量均能反映植物叶片获取营养的能力(Wang et al., 2014)。目前, 众多研究表明温带森林树木叶片从树冠下部到树冠上部, 比叶面积减少而干物质含量呈递增趋势(Asner et al., 2017; Coble & Cavaleri, 2017; Jiang et al., 2017; 田俊霞等, 2018; 荀彦涵等, 2020)。生长在乔木不同冠层的苔藓叶功能性状存在显著差异(Waite & Sack, 2010)。基于草本层、灌木层和林冠层植物功能性状的研究结果表明在森林的不同垂直层次下, 植物表现出明显的性状分异(Vivek & Parthasarathy, 2018; Yang et al., 2018; 李艳朋, 2021)。另外, 乔木层以下的植物叶功能性状随垂直高度变化也表现出一定的适应性, 例如Digrado等(2020)研究发现豇豆(Vigna unguiculata)光合作用和水分利用效率存在冠层差异。然而, 冠层高度引起叶功能性状变异的研究多集中于乔灌木, 对于个体较小的草本层植物, 叶片垂直高度对功能性状的影响是否具有相同结论仍有待探究。
以往研究表明土壤呼吸、养分循环受森林小气候影响(Smith & Johnson, 2004; Ewers & Banks-Leite, 2013)。森林冠层覆盖度的大小决定了土壤湿度的变化, 从而影响蕨类植物的物种组成(Ewers & Banks-Leite, 2013; Cicuzza & Mammides, 2022)。全球尺度下的研究表明, 土壤pH对植物光合性状存在显著影响(Maire et al., 2015; Wang et al., 2017; Cornwell et al., 2018)。然而, 局域尺度下土壤pH是否具有相同结论仍有待考证。土生蕨类植物的根系能够直接从土壤中汲取水分及养分, 土壤资源有限时, 叶片具有更保守的策略(Delpiano et al., 2020)。土壤养分、土壤水分和降水量的结合影响陆地蕨类的功能性状组成(Viana & Dalling, 2022)。综上所述, 土壤作为环境因子, 对蕨类植物叶功能性状变异具有显著影响, 但土壤因子对不同垂直高度叶功能性状变异的影响缺少相关研究。
蕨类从大约4亿年前进化而来(Lehtonen et al., 2017; Morris et al., 2018), 现存的蕨类植物构成了地球上最大的孢子繁殖的维管植物群, 具有重要的研究价值。对蕨类植物性状和生态策略变化的了解, 有助于理解植物进化。近年来, 关于蕨类叶功能性状的研究日益增多。众多研究表明, 从种子植物中发现的叶经济谱同样适用于蕨类植物(Yang et al., 2018; Lin et al., 2020; Li et al., 2022)。相比种子植物, 蕨类植物比叶质量和单位面积氮含量较低, 但单位质量氮含量较高, 这一结果与草本植物模式(Reich et al., 1997; Díaz et al., 2016; Jin et al., 2021)一致。基于不同生境蕨类植物的叶功能性状研究表明蕨类叶性状与生境的相关性十分显著(宋娟等, 2013; 卢艺苗, 2021)。林下草本植物垂直高度变化引起的资源分配也存在差异(Brock et al., 2018), 然而, 关于蕨类垂直高度变化对叶功能性状的影响仍未见报道。
本研究在我国东北东部山区的地带性植被阔叶红松(Pinus koraiensis)林内开展, 以常见的3种大型蕨类(图1)为研究对象, 分别为粗茎鳞毛蕨(Dryopteris crassirhizoma)、东北蹄盖蕨(Athyrium brevifrons)和荚果蕨(Matteuccia struthiopteris)。3种蕨类均属多年生草本, 地上部分每年抽出新叶, 叶片生长周期短。测定不同垂直高度叶片的形态、生理和化学性状, 包括比叶面积、叶干物质含量、净光合速率、瞬时水分利用效率、叶氮和叶磷含量共6种, 并记录个体所处光环境和土壤因子, 旨在研究以下内容: (1)揭示阔叶红松林3种典型蕨类叶功能性状的垂直变异规律, 并分析种间差异; (2)分析叶功能性状之间的相关关系, 并探究叶片垂直高度对性状间相关关系的影响; (3)分析光环境和土壤环境因子对不同垂直高度叶功能性状变异的影响。
图1
图1
阔叶红松林3种典型蕨类。L,下层; M, 中层; U, 上层。
Fig. 1
Three typical ferns in mixed broadleaved-Korean pine forest. L, lower layer; M, middle layer; U, upper layer. AN, available nitrogen; AP, available phosphorus; SWC, soil water content; TN, total nitrogen; TP, total phosphorus.
1 材料和方法
1.1 研究区域概况
凉水国家级自然保护区(128.79°-128.96° E, 47.11°-47.27° N)隶属于黑龙江省伊春市, 属温带大陆性季风气候, 夏季高温多雨, 冬季寒冷干燥, 年平均气温-0.3 ℃, 年降水量676 mm, 年蒸发量855 mm, 地带性植被为阔叶红松林, 森林覆被率为96%, 草本层有葶苈(Draba nemorosa)、东北点地梅(Androsace filiformis)、东北穗花(Pseudolysimachion rotundum subsp. subintegrum)、狼尾草(Pennisetum alopecuroides)等一、二年生草本植物和蚊子草(Filipendula palmata)、宽叶荨麻(Urtica laetevirens)、东北羊角芹(Aegopodium alpestre)、粗茎鳞毛蕨等多年生草本植物, 地带性土壤为暗棕壤, 土壤pH介于4.55-6.32, 呈弱酸性。
1.2 样品采集
2021年8月, 在保护区阔叶红松林内坡向相同、坡度相似、平均冠层开放度为24.3%左右的林内生境下, 对满足叶片垂直高度在75-85 cm范围内的粗茎鳞毛蕨、东北蹄盖蕨和荚果蕨随机采样, 每种蕨类带土挖取10株长势良好, 发育成熟的个体, 每株个体距离保证在5 m以上, 以减少空间自相关性的影响。以最贴近地面的羽叶至植株最高处的垂直高度作为个体叶片垂直高度, 按垂直高度将叶片均等平分为上、中、下3个层次, 并在各层中间位置各取1片健康且完全展开的叶片用于光合及形态性状测量, 其余叶片采集后用自封袋保存带回实验室。
1.3 性状测量
1.3.1 叶功能性状测定
在天气晴朗的上午9:00-11:00, 用LI-6800便携式光合仪(LI-COR, Lincoln, USA)测定叶光合参数。设置叶室参数为: 光强1 500 μmol·m-2·s-1, CO2浓度400 μmol·mol-1, 叶室温度25 ℃, 相对湿度65%。每株植物每层各取1片完整样叶, 在设定光强下诱导 5 min后, 测定其净光合速率(µmol·m-2·s-1)和蒸腾速率(mol·m-2·s-1), 计算得到瞬时水分利用效率(µmol·mol-1), 计算公式为: 瞬时水分利用效率=净光合速率/蒸腾速率。
光合性状测定后将叶片保存带回实验室用于形态性状测量。利用Canon LiDE 400扫描仪测量叶面积; 利用电子天平(精度0.000 1 g)称量样叶鲜质量后放入65 ℃烘箱烘干至恒质量, 之后用电子天平测定叶干质量。叶面积与叶干质量的比值即为比叶面积(cm2·g-1), 叶干质量与叶鲜质量的比值即为干物质含量(g·g-1)。
其余叶片在65 ℃烘箱内烘干至恒质量后研磨至粉状, 过100目筛, 消煮后, 采用AQ400自动间断化学分析仪(SEAL Analytical, Mequon, USA)测定叶片全氮含量(mg·g-1)和全磷含量(mg·g-1)。
1.3.2 光环境及土壤因子测定
采用半球摄影法(digital hemispherical photography, DHP)测定光环境因子, 使用Gap光分析仪软件(GLA)对鱼眼照片进行分析, 以冠层开放度(CO)作为光环境指标。每株个体周围选取3个点采集鲜土样品并混合。鲜土去除石块、细根后装入铝盒并做好标记, 105 ℃下烘干72 h后测定土壤水分含量(SWC); 剩余鲜土风干后磨粉过100目筛, 消煮后, 采用AQ400自动间断化学分析仪测定土壤全氮(TN)含量和全磷(TP)含量。采用碱解扩散法测定有效氮(AN)含量, 钼锑抗比色法测定有效磷(AP)含量。采用电位法测定土壤pH。本研究3种蕨类样本所处生境的环境因子概况见表1。
表1 3种蕨类所处生境的环境因子概况(平均值±标准误)
Table 1
环境因子 Environmental factor | 粗茎鳞毛蕨 Dryopteris crassirhizoma | 东北蹄盖蕨 Athyrium brevifrons | 荚果蕨 Matteuccia struthiopteris |
---|---|---|---|
冠层开放度 Canopy openness (%) | 24.29 ± 1.04a | 24.90 ± 0.66a | 23.68 ± 2.45a |
土壤水分含量 Soil water content | 0.90 ± 0.09a | 0.87 ± 0.17a | 0.99 ± 0.12a |
土壤全氮含量 Soil total nitrogen content (mg·g-1) | 9.32 ± 0.74a | 6.57 ± 0.91b | 9.06 ± 0.64ab |
土壤全磷含量 Soil total phosphorus content (mg·g-1) | 0.82 ± 0.08a | 0.89 ± 0.16a | 0.88 ± 0.10a |
土壤pH Soil pH | 5.67 ± 0.11ab | 5.31 ± 0.10b | 5.76 ± 0.11a |
土壤有效氮含量 Soil available nitrogen content (mg·g-1) | 0.59 ± 0.05a | 0.50 ± 0.06a | 0.59 ± 0.03a |
土壤有效磷含量 Soil available phosphorus content (mg·g-1) | 0.02 ± 0.00a | 0.01 ± 0.00a | 0.02 ± 0.00a |
同行不同小写字母表示不同蕨类间差异显著(p < 0.05)。
Different lowercase letters in the same row indicate significant difference between ferns (p < 0.05).
1.4 数据分析
首先对数据进行对数转换以保证数据正态化。在叶片垂直高度和种间水平, 分别对叶性状做单因素方差分析并进行Tukey检验, 显著性水平设置为α = 0.05; 采用Pearson相关性分析比较不同高度性状间的相关性; 对有显著相关关系的性状采用标准化主轴估计(SMA)法检验不同垂直高度蕨类叶功能性状间的相关关系是否存在显著差异; 若不同高度叶功能性状相关关系存在共同斜率, 则表明功能性状间关系在高度上不存在显著差异; 如不存在显著差异, 再对其截距进行差异性检验。性状间的关系用方程y = bxa来描述, 线性转换为lgy = lgb + algx, x、y分别表示两个功能性状, lgb表示截距, a表示斜率。当|a| = 1时, x、y为等速变化; 当|a|与1有显著差异时, x、y为异速变化。采用冗余分析法(RDA)分析环境因子对功能性状的影响。以上统计分析用到SPSS 15.0、R语言的smatr函数和Canoco 5软件。
2 结果
2.1 叶功能性状的垂直变异
图2
图2
阔叶红松林3种典型蕨类叶功能性状的垂直变异。A.bre, 东北蹄盖蕨; D.cra, 粗茎鳞毛蕨; M.str, 荚果蕨。LDMC, 叶干物质含量; LNC, 叶氮含量; LPC, 叶磷含量; Pn, 净光合速率; SLA, 比叶面积; WUEi, 瞬时水分利用效率。不同大写字母表示种间差异显著(p < 0.05), 不同小写字母表示种内差异显著(p < 0.05)。黑色虚线为平均值。
Fig. 2
Differences in leaf functional traits of three typical ferns in mixed broadleaved-Korean pine forest. A.bre, Athyrium brevifrons; D.cra, Dryopteris crassirhizoma; M.str, Matteuccia struthiopteris. LDMC, leaf dry matter content; LNC, leaf nitrogen content; LPC, leaf phosphorus content; Pn, leaf net photosynthetic rate; SLA, specific leaf area; WUEi, instantaneous water use efficiency. Different uppercase letters indicate significant differences between species (p < 0.05), and different lowercase letters indicate significant intraspecific differences (p < 0.05). The black dashed line represents the average value.
2.2 不同垂直高度叶功能性状的相关性
图3
图3
阔叶红松林3种典型蕨类不同垂直高度叶功能性状的相关性。Corr, 相关系数; L, 下层; M, 中层; U, 上层。A, 净光合速率; LDMC, 叶干物质含量; LNC, 叶氮含量; LPC, 叶磷含量; SLA, 比叶面积; WUEi, 瞬时水分利用效率。*, p < 0.05; **, p < 0.01; ***, p < 0.001。
Fig. 3
Correlation of leaf functional traits at different vertical heights in three typical ferns in mixed broadleaved-Korean pine forest. Corr, correlation coefficient; L, lower layer; M, middle layer; U, upper layer. A, leaf net photosynthetic rate; LDMC, leaf dry matter content; LNC, leaf nitrogen content; LPC, leaf phosphorus content; SLA, specific leaf area; WUEi, instantaneous water use efficiency. *, p < 0.05; **, p < 0.01; ***, p < 0.001.
不同蕨类的性状相关关系并无较大差异(附录I)。不同垂直高度下, 叶功能性状的相关关系在斜率上均未表现出显著差异(图4)。叶氮含量与叶干物质含量、叶干物质含量与比叶面积在各垂直高度均呈负相关关系。叶干物质含量一定时, 叶氮含量表现为上层>中层>下层(图4D); 叶干物质含量与比叶面积在不同垂直高度截距无显著差异(图4E); 叶氮含量与比叶面积、瞬时水分利用效率与净光合速率则呈显著正相关关系, 且在不同垂直高度存在相同斜率和截距(图4F、4H)。另外, 一些性状仅在某些高度达到显著的相关关系。例如, 净光合速率与比叶面积仅在上层呈负相关关系(图4A); 净光合速率与叶氮含量仅在下层呈负相关关系(图4B); 净光合速率与叶干物质含量在上、下层呈正相关关系(图4C); 叶磷含量与叶干物质含量在上、下层呈显著负相关关系(图4G)。此外,和叶氮含量与叶干物质含量的匀速生长不同, 其他性状之间均表现为异速生长(附录II)。
图4
图4
阔叶红松林3种典型蕨类的叶功能性状相关关系在不同高度间的差异。A, 净光合速率; LDMC, 叶干物质含量; LNC, 叶氮含量; LPC, 叶磷含量; SLA, 比叶面积; WUEi, 瞬时水分利用效率。Common slope, 共同斜率; Slope, 斜率。图中彩色实线表示性状间相关关系显著, 黑色实线表示3条实线重合。
Fig. 4
Differences in leaf trait correlations among different heights for three typical ferns in mixed broadleaved-Korean pine forest. A, leaf net photosynthetic rate; LDMC, leaf dry matter content; LNC, leaf nitrogen content; LPC, leaf phosphorus content; SLA, specific leaf area; WUEi, instantaneous water use efficiency. The colored solid lines in the figure indicate that the correlations between traits were significant, the black solid line indicates that the three solid lines overlap.
2.3 叶功能性状与环境因子之间的冗余分析
光环境和土壤因子对上、中、下层叶功能性状变异的解释率依次为27.9%、23.6%和25.6% (附录III)。对于上层(图5A), 土壤全氮含量、土壤有效氮含量和土壤有效磷含量与比叶面积、叶氮含量呈显著负相关关系, 与净光合速率、叶干物质含量、瞬时水分利用效率呈正相关关系(图5A)。对于中层, 土壤全磷含量、有效磷含量和土壤水分含量与叶氮含量、比叶面积的夹角大于90°, 为负相关关系, 与净光合速率、叶干物质含量呈正相关关系; 土壤pH与瞬时水分利用效率存在正相关关系, 与叶磷含量为负相关关系(图5B)。对于下层, 土壤全氮含量、土壤有效磷含量和冠层开放度与净光合速率、瞬时水分利用效率呈正相关关系, 土壤pH与比叶面积、叶氮含量呈显著正相关关系(图5C)。
图5
图5
阔叶红松林3种典型蕨类不同垂直高度下叶功能性状与环境因子的冗余分析(RDA)排序。A, 上层。B, 中层。C, 下层。A, 净光合速率; AN, 土壤有效氮含量; AP, 土壤有效磷含量; CO, 冠层开放度; LDMC, 叶干物质含量; LNC, 叶氮含量; LPC, 叶磷含量; pH, 土壤pH; SLA, 比叶面积; SWC, 土壤水分含量; TN, 土壤全氮含量; TP, 土壤全磷含量; WUEi, 瞬时水分利用效率。
Fig. 5
Redundancy analysis (RDA) ranking of leaf functional traits and environmental factors in three typical ferns in mixed broadleaved-Korean pine forest. A, Lower layer; B, Middle layer; C, Upper layer. A, leaf net photosynthetic rate; AN, soil available nitrogen content; AP, soil available phosphorus content; CO, canopy openness; LDMC, leaf dry matter content; LNC, leaf nitrogen content; LPC, leaf phosphorus content; pH, soil pH; SLA, specific leaf area; SWC, soil water content; TN, soil total nitrogen content; TP, soil total phosphorus content; WUEi, instantaneous water use efficiency.
3 讨论
3.1 叶功能性状的垂直变化规律
本研究表明, 相比东北蹄盖蕨和荚果蕨, 粗茎鳞毛蕨具有小比叶面积, 高叶干物质含量, 叶氮含量、叶磷含量显著偏低的特点, 但其净光合速率和瞬时水分利用效率则高于其他2种蕨类; 荚果蕨的叶氮含量较高但净光合速率偏低, 这与以往净光合速率与叶氮含量有相同变化趋势的结果(Tosens et al., 2016)不一致。植物叶功能性状随垂直高度的增加表现出明显变化趋势(Leal & Thomas, 2003)。以往研究表明, 光照和水分对植物叶功能性状垂直梯度变异存在重要影响(田俊霞等, 2018; 罗恬等, 2022)。蕨类比叶面积随高度变化无显著差异, 叶干物质含量随高度从高到低递减; 东北蹄盖蕨和荚果蕨净光合速率随高度从高到低呈递减趋势。对处于优势层的高大乔木来说, 上层叶片由于高度优势, 在竞争中率先获取大部分光照; 为避免光强过大、失水过多, 上层叶片会采取减少受光面积、增加角质层厚度、关闭气孔等措施; 蕨类由于长期生活在林下弱光环境中, 上层获得光照优势, 光合作用积累的产物较多, 但不需要减少受光面以避免强光照射, 表现为净光合速率较强, 叶干物质含量高, 比叶面积无显著差异; 下层叶片长期处于光照更弱的环境, 限制叶片生长, 因而表现为净光合速率和叶干物质含量较中上层更低(宋娟等, 2013; Coble et al., 2017)。粗茎鳞毛蕨和东北蹄盖蕨不同高度叶片的瞬时水分利用效率无显著差异, 荚果蕨叶片瞬时水分利用效率随高度升高呈先增后减的趋势, 一定程度反映不同高度叶片蒸腾速率的变化及对干旱环境的适应能力, 即上层叶片的蒸腾速率显著高于中下层; 中层较上下层而言, 具有更强的耐旱能力。不同垂直高度下, 荚果蕨的叶氮含量和粗茎鳞毛蕨的叶磷含量有较大差异, 反映不同垂直高度不同种类的植物叶片对营养资源的利用能力有较大差别, 氮和磷可能分别为荚果蕨和粗茎鳞毛蕨叶片生长的限制元素。
3.2 垂直高度对叶片性状相关关系的影响
本研究结果表明, 蕨类净光合速率随叶干物质含量的增大而增大, 这与罗达等(2021)对榛(Corylus heterophylla)种质资源叶功能性状的研究结果一致;叶干物质含量与比叶面积呈负相关关系, 这与许多研究结果(de la Riva et al., 2016; 刘贵峰等, 2017)一致。垂直高度下净光合速率与叶干物质含量、叶干物质含量与比叶面积相关关系的斜率与截距无显著差异, 说明林下的弱光环境对蕨类各层叶功能性状间的生物量分配及资源利用的影响不显著。蕨类作为林下草本植物, 被林内高大乔木和灌木的遮蔽, 接受到的光照从上到下依次减少, 不同垂直高度的叶片充分利用弱光进行光合作用, 光合速率随接受光照的强弱变化, 即上层叶片净光合速率高于下层, 从而造成生长位置靠下的叶片具有更低的叶干物质含量。比叶面积减小, 即叶干物质含量增大, 主要与叶肉细胞层厚度增大, 叶片变厚, 叶片密度显著增强, 细胞内含物显著上升有关(刘明秀和梁国鲁, 2016)。
氮是构成叶绿素的重要物质, 在调节光合作用、资源运输和繁殖等方面发挥重要作用(Allison, 2002; Lovelock et al., 2004)。以往研究表明, 单位面积光合速率、单位质量光合速率与单位质量氮含量均呈高度正相关关系(朱军涛等, 2010; Funk et al., 2020; Martin & Isaac, 2021), 然而, 本研究结果与之相反。有学者认为光合能力的差异由比叶质量和氮含量的变化引起, 比叶质量通过影响氮含量进而影响光合速率, 即光合能力主要受比叶质量影响(Le Roux et al., 2001; Walcroft et al., 2002)。通常高比叶质量反映叶片低氮含量, 这与本研究中叶氮含量与比叶面积呈正相关关系的结果一致; 叶氮含量的增加反而引起净光合速率的下降, 这与以往研究结果(朱军涛等, 2010; Khan et al., 2022)相反。Westoby等(2013)提出净光合速率与比叶质量在物种间没有太大的相关性; 在高比叶质量的叶片中, 大量氮可用于非光合成分合成, 从而导致较低的光合速率(Bucher et al., 2021), 本研究结果与之一致。可见, 区域环境下比叶质量是否为蕨类光合能力的主要衡量参数仍值得探究。本研究中虽然将林下蕨类叶片按垂直高度分为3层, 但整体而言, 其生长环境长期荫蔽, 接受的光照主要来源为林隙透光和地面散射, 上层叶片较下层而言受光条件好, 可以更好地利用光照进行光合作用, 净光合速率更高, 因而净光合速率与比叶面积表现为负相关关系。养分含量越高, 植物光合能力和呼吸消耗越强, 高效的代谢有利于植物适应环境(Cornelissen et al., 2003; Wright et al., 2004)。本研究中, 叶氮、磷含量与叶干物质含量均表现为负相关关系, 且各高度相关关系斜率相同, 叶氮含量与叶干物质含量截距表现为上层>中层>下层; 叶磷含量与叶干物质含量截距为上层>下层, 与以往研究结果(荀彦涵等, 2020)一致。受光照条件影响, 叶片内有机物积累随高度升高逐渐增多, 叶干物质含量自下而上依次升高, 叶片内氮、磷含量呈上升趋势。瞬时水分利用效率与净光合速率在不同垂直高度的正相关关系并无差别, 不同垂直高度的光照变化不足以引起叶片耐旱机制的分异。
3.3 光环境和土壤因子对叶功能性状的影响
本研究表明, 随叶片垂直高度的增加, 冠层开放度对叶功能性状的解释率逐渐上升。叶功能性状受光照条件的影响(Wang et al., 2021; Westerband et al., 2021)。与乔、灌木相比, 由于树冠的遮阴, 蕨类的光照条件较差, 光强度也较弱。在阴蔽环境下, 高度较高的叶片受光照强度更大, 促进蕨类叶片光合、蒸腾、呼吸等生理活动, 使叶功能性状向适应环境的方向变化。本研究区土壤呈弱酸性, 该条件下植物对氮的利用率降低(Jenny, 1941), 因而叶片氮含量与土壤总氮含量、土壤有效氮含量在不同垂直高度均表现为负相关关系, 这与以往研究结论(Santiago et al., 2005; Ordoñez et al., 2009)一致。另外, 土壤局部氮含量的变化并不能有效地反映植物可利用氮含量的变化(Burton et al., 2017)。本研究仅下层叶片的叶氮含量受土壤pH影响较大, 地表枯落物分解对土壤pH的调节影响近地面叶片氮的吸收利用。本研究中叶片和土壤磷含量变化具有一致性, 土壤有效磷含量对不同垂直高度叶功能性状变异的解释率最高, 与净光合速率均表现为正相关关系。叶片中磷积累越快, 生长越快(Guo et al., 2020), 3种蕨类属多年生草本, 地上部分每年抽出新叶, 叶片生长周期短, 需要从土壤中吸收大量可用的磷投入叶片以维持自身生长发育。此外, 本研究中蕨类生长环境并无较大差异(表1), 植物叶功能性状的变异受多重环境条件的影响, 温度、湿度、风力等其他环境因素对蕨类叶功能性状的影响仍有待探究。
4 结论
不同种的蕨类叶功能性状存在一定差异, 且一定程度上, 在不同垂直高度下呈现相同的变化趋势。除比叶面积外, 蕨类叶片的其他功能性状随垂直高度的增加变化明显; 叶功能性状间的相关关系在不同冠层位置存在共同斜率, 垂直高度并未引起性状间相关关系的明显变化, 即在个体内叶片变化速率是恒定的, 但不同高度叶片的资源获取及利用能力有所差异。各环境因子对不同垂直高度叶功能性状的影响方向和程度存在差异, 其中影响各层叶功能性状变异的主要环境因子为土壤有效磷含量和土壤pH。除光环境和土壤因子外, 其他环境因素对蕨类叶功能性状的影响还需进一步探究。以叶功能性状为切入点探究不同垂直高度下蕨类叶功能性状的变异及适应机制, 反映出蕨类叶片为适应环境变化所形成的资源利用策略, 对深入了解植物的适应进化具有重要意义。
附录
附录I 阔叶红松林3种典型蕨类不同高度叶功能性状相关关系在种间的差异
Supplement I Differences in leaf trait correlations among different heights for three typical ferns in mixed broadleaved- Korean pine forest
附录II 阔叶红松林3种典型蕨类不同垂直高度下叶功能性状的标准主轴分析结果
Supplement II Standardized major axis analysis results of leaf functional traits at different vertical heights in mixed broadleaved-Korean pine forest
附录III 阔叶红松林3种典型蕨类不同垂直高度环境因子对叶功能性状的解释率
Supplement III Interpretation of leaf functional traits by environmental factors at different vertical heights in mixed broadleaved-Korean pine forest
致谢
感谢黑龙江凉水国家级自然保护区管理局在样品采集过程中给予的大力支持。
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There is growing recognition that classifying terrestrial plant species on the basis of their function (into 'functional types') rather than their higher taxonomic identity, is a promising way forward for tackling important ecological questions at the scale of ecosystems, landscapes or biomes. These questions include those on vegetation responses to and vegetation effects on, environmental changes (e.g. changes in climate, atmospheric chemistry, land use or other disturbances). There is also growing consensus about a shortlist of plant traits that should underlie such functional plant classifications, because they have strong predictive power of important ecosystem responses to environmental change and/or they themselves have strong impacts on ecosystem processes. The most favoured traits are those that are also relatively easy and inexpensive to measure for large numbers of plant species. Large international research efforts, promoted by the IGBP–GCTE Programme, are underway to screen predominant plant species in various ecosystems and biomes worldwide for such traits. This paper provides an international methodological protocol aimed at standardising this research effort, based on consensus among a broad group of scientists in this field. It features a practical handbook with step-by-step recipes, with relatively brief information about the ecological context, for 28 functional traits recognised as critical for tackling large-scale ecological questions.
Climate and soils together regulate photosynthetic carbon isotope discrimination within C3 plants worldwide
DOI:10.1111/geb.v27.9 URL [本文引用: 1]
A plant economics spectrum in Mediterranean forests along environmental gradients: Is there coordination among leaf, stem and root traits?
DOI:10.1111/jvs.2016.27.issue-1 URL [本文引用: 1]
Different responses of leaf and root traits to changes in soil nutrient availability do not converge into a community-level plant economics spectrum
DOI:10.1007/s11104-020-04515-2 [本文引用: 1]
The global spectrum of plant form and function
DOI:10.1038/nature16489 [本文引用: 1]
Assessing diversity in canopy architecture, photosynthesis, and water-use efficiency in a cowpea magic population
Optimizing crops to improve light absorption and CO assimilation throughout the canopy is a proposed strategy to increase yield and meet the needs of a growing population by 2050. Globally, the greatest population increase is expected to occur in Sub-Saharan Africa where large yield gaps currently persist; therefore, it is crucial to develop high-yielding crops adapted to this region. In this study, we screened 50 cowpea ( (L.) Walp) genotypes from the multi-parent advanced generation inter-cross (MAGIC) population for canopy architectural traits, canopy photosynthesis, and water-use efficiency using a canopy gas exchange chamber in order to improve our understanding of the relationships among those traits. Canopy architecture contributed to 38.6% of the variance observed in canopy photosynthesis. The results suggest that the light environment within the canopy was a limiting factor for canopy CO assimilation. Traits favoring greater exposure of leaf area to light such as the width of the canopy relative to the total leaf area were associated with greater canopy photosynthesis, especially in canopies with high biomass. Canopy water-use efficiency was highly determined by canopy photosynthetic activity and therefore canopy architecture, which indicates that optimizing the canopy will also contribute to improving canopy water-use efficiency. We discuss different breeding strategies for future programs aimed at the improvement of cowpea yield for the Sub-Saharan African region. We show that breeding for high biomass will not optimize canopy CO assimilation and suggest that selection should include multiple canopy traits to improve light penetration.© 2020 The Authors. Food and Energy Security published by John Wiley & Sons Ltd on behalf of Association of Applied Biologists.
Fragmentation impairs the microclimate buffering effect of tropical forests
Leaf traits and performance vary with plant age and water availability in Artemisia californica
DOI:10.1093/aob/mcaa106
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Leaf functional traits are strongly tied to growth strategies and ecological processes across species, but few efforts have linked intraspecific trait variation to performance across ontogenetic and environmental gradients. Plants are believed to shift towards more resource-conservative traits in stressful environments and as they age. However, uncertainty as to how intraspecific trait variation aligns with plant age and performance in the context of environmental variation may limit our ability to use traits to infer ecological processes at larger scales.
The community-level scaling relationship between leaf nitrogen and phosphorus changes with plant growth, climate and nutrient limitation
DOI:10.1111/jec.v108.4 URL [本文引用: 1]
Analysis of photosynthetic characteristics of Pinus koraiensis with different slope orientations
不同坡向红松光合特性分析
Coordination and determinants of leaf community economics spectrum for canopy trees and shrubs in a temperate forest in northeastern China
Functional traits: adaption of ferns in forest
DOI:10.1111/jse.12669
[本文引用: 1]
Ferns that evolved from 400 million years ago show various functional traits and ecological strategies in extant species, over 80% of which belong to the youngest order Polypodiales. How the functional traits and strategies of ferns have changed during their evolutionary history remains unexplored. Here, we measured functional traits that are sensitive to environmental light and water availability in 345 fern species across fern phylogeny, and reconstructed their evolutionary histories. We found that ferns, mainly Polypodiales, have developed diversified functional traits in response to forest environments. Terrestrial species, especially Thelypteridaceae and Athyriaceae in eupolypods II, since the late Jurassic period, have shown decreased leaf mass per area (LMA) and area-based leaf nitrogen (N<sub>area</sub>), but increased mass-based leaf nitrogen (N<sub>mass</sub>) compared with early-derived polypods. Epiphytic species, mainly those in Polypodiaceae, have shown reductions in N<sub>mass</sub> and individual leaf area (Area) since the late Cretaceous period. The adaption of functional traits of Polypodiales to forest environment may have played a crucial role in fern radiation since the late Jurassic period. Integrative analysis of functional traits, especially numerical ones, may shed new light on plant evolution.
TRY—A global database of plant traits
Leaf traits and leaf nitrogen shift photosynthesis adaptive strategies among functional groups and diverse biomes
Photosynthetic light acclimation in peach leaves: importance of changes in mass:area ratio, nitrogen concentration, and leaf nitrogen partitioning
DOI:10.1093/treephys/21.6.377
PMID:11282577
[本文引用: 1]
Photosynthetic light acclimation of leaves can result from (i) changes in mass-based leaf nitrogen concentration, Nm, (ii) changes in leaf mass:area ratio, Ma, and (iii) partitioning of total leaf nitrogen among different pools of the photosynthetic machinery. We studied variations in Nm and Ma within the crowns of two peach (Prunus persica L. Batsch) trees grown in an orchard in Portugal, and one peach tree grown in an orchard in France. Each crown was digitized and a 3-D radiation transfer model was used to quantify the intra-crown variations in time-integrated leaf irradiance, <PARi>. Nitrogen concentration, leaf mass:area ratio, chlorophyll concentration, and photosynthetic capacity were also measured on leaves sampled on five additional peach trees in the orchard in Portugal. The data were used to compute the coefficients of leaf nitrogen partitioning among carboxylation, bioenergetics, and light harvesting pools. Leaf mass:area ratio and area-based leaf nitrogen concentration, Na, were nonlinearly related to <PARi>, and photosynthetic capacity was linearly related to Na. Photosynthetic light acclimation resulted mainly from changes in Ma and leaf nitrogen partitioning, and to a lesser extent from changes in Nm. This behavior contrasts with photosynthetic light acclimation observed in other tree species like walnut (Juglans regia L.) in which acclimation results primarily from changes in Ma.
Vertical gradients and tree-to-tree variation in shoot morphology and foliar nitrogen in an old-growth Pinus strobus stand
DOI:10.1139/x03-064
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Acclimation to ambient light has been assumed to be the principal cause of vertical gradients in leaf nitrogen, foliar morphology, and related traits in forest canopies. We examined the relative role of crown exposure, damage, and reproduction as correlates of vertical gradients in shoot and needle morphology, anatomy, and chemistry in a ~120-year-old stand of Pinus strobus L. (eastern white pine) in central Ontario. Internodes at the top of trees were longer, wider, and produced more fascicles, but had lower fascicle survivorship than lower-canopy shoots. Needles on upper-canopy shoots also had higher allocation to resin ducts than mid- or lower-canopy needles, contained less nitrogen on a mass basis, and showed a higher leaf mass per area, C/N ratio, and chlorophyll a/b ratio. Tree-to-tree variation among the 26 trees measured traits was high, owing, in part, to differences in crown damage and reproductive status. Crown damage was associated with reduced leaf mass per area and other traits associated with shade acclimation, while high cone production was associated with reduced leaf nitrogen and chlorophyll in the upper canopy. Our results suggest that factors other than light acclimation play an important role in determining vertical gradients in foliar morphology, nitrogen, and leaf production in forest canopies.
Environmentally driven extinction and opportunistic origination explain fern diversification patterns
Combining palaeontological and neontological data offers a unique opportunity to investigate the relative roles of biotic and abiotic controls of species diversification, and the importance of origination versus extinction in driving evolutionary dynamics. Ferns comprise a major terrestrial plant radiation with an extensive evolutionary history providing a wealth of modern and fossil data for modelling environmental drivers of diversification. Here we develop a novel Bayesian model to simultaneously estimate correlations between diversification dynamics and multiple environmental trajectories. We estimate the impact of different factors on fern diversification over the past 400 million years by analysing a comprehensive dataset of fossil occurrences and complement these findings by analysing a large molecular phylogeny. We show that origination and extinction rates are governed by fundamentally different processes: originations depend on within-group diversity but are largely unaffected by environmental changes, whereas extinctions are strongly affected by external factors such as climate and geology. Our results indicate that the prime driver of fern diversity dynamics is environmentally driven extinction, with origination being an opportunistic response to diminishing ecospace occupancy.
Canopy characteristics in gaps and its relationship with seedlings and saplings in a spruce-fir forest in the Changbai Mountain area of northeastern China
长白山云冷杉林林隙冠层特征及与幼苗幼树的关系
The fern economics spectrum is unaffected by the environment
Relationship between variation of plant functional traits and individual growth at different vertical layers in a subtropical evergreen broad-leaved forest of Dinghushan
DOI:10.17520/biods.2021205 URL [本文引用: 1]
鼎湖山南亚热带常绿阔叶林植物功能性状变异与不同垂直层次个体生长的关联
DOI:10.17520/biods.2021205
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建立植物功能性状与群落动态之间的关联是功能生态学的核心问题之一。本文基于鼎湖山1.44 ha塔吊样地的两次调查数据, 通过采集样地内所有4,142株个体的6种植物功能性状, 对比分析了个体水平植物功能性状和物种水平功能性状均值对不同垂直层次(灌木层、亚冠层和林冠层)个体生长的影响差异。首先, 分析了不同垂直层次下各植物功能性状的变化趋势; 其次, 计算了不同垂直层次下各植物功能性状的种内和种间变异水平; 最后, 运用结构方程模型探讨了植物功能性状、光竞争以及地下竞争对不同垂直层次树木生长的影响。结果表明: (1)不同垂直层次下的植物功能性状表现出明显的分异, 由灌木层至林冠层, 叶面积、比叶面积和能量供求关系指数显著降低, 而叶片厚度和叶片干物质含量显著升高; (2)不同垂直层次下植物功能性状的种间变异均大于种内变异, 且林冠层的种内功能性状变异均大于灌木层和亚冠层; (3)基于个体水平植物功能性状的结构方程模型较物种水平功能性状均值对生长具有更高的解释程度, 且个体水平植物功能性状的引入更有利于提高对灌木层个体生长的预测能力; (4)光竞争和地下竞争主要通过影响功能性状间接影响植物生长。由灌木层至林冠层, 同种间的相互作用逐渐减弱, 异种间的相互作用逐渐增强。综上, 将个体水平植物功能性状纳入分析有助于更好地理解群落的结构和动态。
A plant economics spectrum of litter decomposition among coexisting fern species in a sub-tropical forest
DOI:10.1093/aob/mcz166
PMID:31633171
[本文引用: 1]
The plant economics spectrum theory provides a useful framework to examine plant strategies by integrating the co-ordination of plant functional traits along a resource acquisition-conservation trade-off axis. Empirical evidence for this theory has been widely observed for seed plants (Spermatophyta). However, whether this theory can be applied to ferns (Pteridophyta), a ubiquitous and ancient group of vascular plants, has rarely been evaluated so far.We measured 11 pairs of plant functional traits on leaves and fine roots (diameter <2 mm) on 12 coexisting fern species in a sub-tropical forest. Litterbags of leaves and roots were placed in situ and exposed for 586 d to measure decomposition rates. The variation of traits across species and the co-ordination among traits within and between plant organs were analysed. Finally, the influence of the traits on decomposition rates were explored.Most leaf and root traits displayed high cross-species variation, and were aligned along a major resource acquisition-conservation trade-off axis. Many fern traits co-varied between leaves and fine roots, suggesting co-ordinated responses between above- and below-ground organs. Decomposition rates of leaves were significantly higher than those of fine roots, but they were significantly and positively correlated. Finally, our results highlight that the decomposition of both leaves and roots was relatively well predicted by the leaf and root economics spectra.Our results support the existence of an acquisition-conservation trade-off axis within ferns and indicate that traits have important 'afterlife' effects on fern litter decomposition. We conclude that the plant economics spectrum theory that is commonly observed across seed plants can be applied to ferns species, thereby extending the generality of this theory to this ancient plant lineage in our study site. Our study further suggests that the evolutionary and ecological basis for the relationships among key economics traits appears to be similar between ferns and seed plants. Future studies involving larger data sets will be required to confirm these findings across different biomes at larger spatial scales.© The Author(s) 2019. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.
Leaf traits of dominant plants of main forest communities in Daqinggou Nature Reserve
大青沟自然保护区主要森林群落优势种的叶性状
Research progress on leaf mass per area
DOI:10.17521/cjpe.2015.0428 URL [本文引用: 1]
植物比叶质量研究进展
DOI:10.17521/cjpe.2015.0428
[本文引用: 1]
比叶质量(LMA)是叶片经济型谱中最基础的叶功能性状, 也是重要的复合型结构参数。LMA不仅与植物的许多生理反应密切相关, 而且能定量测定光合产物在单位叶面积的投入, 因而被认为是反映植物生态策略的重要指标。目前, 有关LMA的深入研究已在植物生态学、农学、林学、植物生理学领域全面展开。该文系统阐述了LMA在叶片整体、组织、细胞三个水平的结构解析和计算方法; 重点分析了LMA对光合作用的影响; 讨论了LMA的内在遗传差异以及外部的环境胁迫因子(温度、水分、光照)对LMA的影响, 以期梳理比叶质量研究的思路、策略和方法, 为今后的研究提供借鉴和参考。
Plant functional traits—Concepts, applications and future directions
植物功能性状研究进展
The effect of nutrient enrichment on growth, photosynthesis and hydraulic conductance of dwarf mangroves in Panamá
DOI:10.1046/j.0269-8463.2004.00805.x
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\n\nDwarf stands of the mangrove Rhizophora mangle L. are extensive in the Caribbean. We fertilized dwarf trees in Almirante Bay, Bocas del Toro Province, north‐eastern Panamá with nitrogen (N) and phosphorus (P) to determine (1) if growth limitations are due to nutrient deficiency; and (2) what morphological and/or physiological factors underlie nutrient limitations to growth.
Variation and correlation of leaf functional traits and photosynthetic characteristics of 38 hazelnut germplasm resources
38个榛种质资源叶功能性状与光合特征变异及其相关性
Impact of canopy vertical height on leaf functional traits in a lower subtropical evergreen broad-leaved forest of Dinghushan
<p id="p00005"><strong>Aims:</strong> Plant functional traits, which reflect the adaptation mechanisms of plants to their environment, are important for ecosystem function. Leaves are the main photosynthetic organs of plants and the primary research object of functional traits. However, there are few studies about the variation of leaf functional traits with the increase of canopy vertical height. Taking a lower subtropical evergreen broad-leaved forest of Dinghushan as example, our study aims to detect the changes of leaf functional traits with the canopy vertical height, and explore the influence of canopy vertical height on leaf functional traits.</p><p id="p00010"><strong>Methods:</strong> Six dominant species from a subtropical evergreen broad-leaved forest in the Dinghushan National Nature Reserve and a total of 10 leaf functional traits of these species at different canopy heights were compared and analyzed in this study. Leaf functional traits include specific leaf area (SLA), chlorophyll relative content (SPAD), leaf dry matter content (LDMC), leaf carbon content (LCC), leaf nitrogen content (LNC), leaf phosphorus content (LPC), leaf N : P (N : P), equivalent water thickness (EWT), nitrogen balance index (NBI), and flavonoids content (FLAV).</p> <p id="p00015"><strong>Results:</strong> The results revealed that: (1) With the increase of canopy vertical height, LDMC, FLAV, EWT, and LPC exhibited an increasing trend, while SLA and NBI exhibited a decreasing trend. There was a significant correlation between leaf traits and vertical height, as exemplified by FLAV and NBI, SLA and EWT with a negative correlation, while LNC and LPC were positively correlated. (2) Leaf traits showed significant discrepancy among different tree species, which might be affected by the plant’s own characteristics and the succession process of species in the community. (3) The intraspecific variation of leaf traits demonstrated different trends along vertical layers. With the increase of canopy vertical height, 67% of species displayed a significant increase in FLAV, while the NBI of 67% of species decreased significantly.</p> <p id="p00020"><strong>Conclusion:</strong> The variation of leaf traits with the increase of canopy vertical height in the subtropical evergreen broad-leaved forest reflects the adaptation of plants to the light environment and the utilization of water resources and energy.</p>
冠层垂直高度对植物叶片功能性状的影响: 以鼎湖山南亚热带常绿阔叶林为例
植物功能性状能够反映植物对环境变化的响应, 从而影响生态系统功能。叶片是植物功能性状的重要研究对象, 但随着树冠垂直高度的增加, 植物叶片功能性状的变化规律仍不清楚。为了更好地解释植物对所处环境的响应机制, 本研究以鼎湖山南亚热带常绿阔叶林6个优势树种为研究对象, 对这些树种不同冠层高度的叶片比叶面积(SLA)、叶绿素相对含量(SPAD)、叶干物质含量(LDMC)、叶片碳含量(LCC)、叶片氮含量(LNC)、叶片磷含量(LPC)、叶片氮磷比(N : P)、等效水厚度(EWT)、氮平衡指数(NBI)、类黄酮含量(FLAV)共10个功能性状指标进行比较分析。结果表明: (1)随树冠垂直高度的增加, 叶干物质含量、类黄酮含量、等效水厚度和叶片磷含量呈增加趋势, 比叶面积和氮平衡指数呈减小趋势; 在冠层垂直方向上, 类黄酮含量与氮平衡指数、比叶面积与等效水厚度呈极显著负相关, 叶片氮含量与叶片磷含量呈极显著正相关。(2)不同树种叶片功能性状差异显著, 可能受植物自身特性和群落树种演替进程的影响。(3)叶片功能性状的种内差异沿垂直层次呈现不同的变化趋势。其中, 67%的树种的类黄酮含量随树冠垂直高度增加而显著增加, 67%的树种的氮平衡指数随树冠垂直高度增加而显著减小。南亚热带常绿阔叶林不同树种叶片功能性状随树冠垂直高度增加产生的变化, 反映了植物对光环境的适应以及对水资源和能量利用的不同策略。
Global effects of soil and climate on leaf photosynthetic traits and rates
DOI:10.1111/geb.2015.24.issue-6 URL [本文引用: 1]
The leaf economics spectrum’s morning coffee: plant size-dependent changes in leaf traits and reproductive onset in a perennial tree crop
DOI:10.1093/aob/mcaa199
PMID:33502446
[本文引用: 1]
Size-dependent changes in plant traits are an important source of intraspecific trait variation. However, there are few studies that have tested if leaf trait co-variation and/or trade-offs follow a within-genotype leaf economics spectrum (LES) related to plant size and reproductive onset. To our knowledge, there are no studies on any plant species that have tested whether or not the shape of a within-genotype LES that describes how traits covary across whole plant sizes, is the same as the shape of a within-genotype LES that represents environmentally driven trait plasticity.We quantified size-dependent variation in eight leaf traits in a single coffee genotype (Coffea arabica var. Caturra) in managed agroecosystems with different environmental conditions (light and fertilization treatments), and evaluated these patterns with respect to reproductive onset. We also evaluated if trait covariation along a within-genotype plant-size LES differed from a within-genotype environmental LES defined with trait data from coffee growing in different environmental conditions.Leaf economics traits related to resource acquisition - maximum photosynthetic rates (A) and mass-based leaf nitrogen (N) concentrations - declined linearly with plant size. Structural traits - leaf mass, leaf thickness, and leaf mass per unit area (LMA) - and leaf area increased with plant size beyond reproductive onset, then declined in larger plants. Three primary LES traits (mass-based A, leaf N and LMA) covaried across a within-genotype plant-size LES, with plants moving towards the 'resource-conserving' end of the LES as they grow larger; in coffee these patterns were nearly identical to a within-genotype environmental LES.Our results demonstrate that a plant-size LES exists within a single genotype. Our findings indicate that in managed agroecosystems where resource availability is high the role of reproductive onset in driving within-genotype trait variability, and the strength of covariation and trade-offs among LES traits, are less pronounced compared with plants in natural systems. The consistency in trait covariation in coffee along both plant-size and environmental LES axes indicates strong constraints on leaf form and function that exist within plant genotypes.© The Author(s) 2021. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.
Plant functional traits, environments and ecosystem functioning
植物功能性状与环境和生态系统功能
DOI:10.17521/cjpe.2007.0019
[本文引用: 1]
植物性状反映了植物对生长环境的响应和适应,将环境、植物个体和生态系统结构、 过程与功能联系起来(所谓的“植物功能性状”)。该文介绍了植物功能性状的分类体系,综述了国内外植物功能性状与气候(包括气温、降水、光照)、地理空间变异(包括地形地 貌、生态梯度、海拔)、营养、干扰(包括火灾、放牧、生物入侵、土地利用)等环境因素,以及与生态系统功能之间关系的研究进展,探讨了全球变化(气候变化和CO<sub>2</sub>浓度升高 ) 对个体和群落植物功能性状的影响。植物功能性状的研究已经取得很多成果,并应用于全球变化、古植被恢复和古气候定量重建、环境监测与评价、生态保护和恢复等研究中,但大尺度、多生境因子下的植物功能性状研究仍有待于加强,同时需要改进性状的测量手段;我国 的植物功能性状研究还需要更加明朗化和系统化。
The timescale of early land plant evolution
A global study of relationships between leaf traits, climate and soil measures of nutrient fertility
DOI:10.1111/geb.2009.18.issue-2 URL [本文引用: 1]
How is light interception efficiency related to shoot structure in tall canopy species?
DOI:10.1007/s00442-017-3926-0
PMID:28801737
[本文引用: 1]
Coexistence of multiple species is a fundamental aspect of plant and forest ecology. Although spatial arrangement of leaves within crowns is an important determinant of light interception and productivity, shoot structure varies considerably among coexisting canopy species. We investigated the relative importance of structural traits in determining the light availability of leaves (I) and light interception efficiency at the current-year shoot level (LIE; the total light interception of leaves divided by shoot biomass) at the top of crowns of 11 canopy species in a cool-temperate forest in Japan. In accordance with Corner's rules, the total mass, stem mass, total mass of leaf laminae, individual leaf area, and stem cross-sectional area of current-year shoot were positively correlated with each other, and branching intensity (the number of current-year shoots per branch unit of 1-m length) was inversely correlated with these traits across species. In contrast, I was correlated not with these traits, but with leaf elevation angle (a ). Moreover, variation in LIE across species was caused by variation in I (thus in a ). Thus, a is a key parameter for the leaf light interception of canopy shoots in this cool-temperate forest. Differences in a across species might be related to different physiological strategies that developed in the high light and water-limited environment of forest canopies. Small variation in the length of current-year shoots among species implies that variations in I and LIE would be important for the coexistence of these canopy species.
Photosynthetic characteristics of two Cylindrospermopsis raciborskii strains differing in their toxicity
DOI:10.1111/jpy.12157
PMID:26988186
[本文引用: 1]
We studied the growth and photosynthetic characteristics of a toxic (CS506) and a nontoxic strain (CS509) of the bloom-forming cyanobacterium Cylindrospermopsis raciborskii grown under identical experimental conditions. When exposed to light-saturating growth conditions (100 μmol photons · m(-2) · s(-1) ), values for maximal photosynthetic capacity (Pmax ) and maximum quantum yield (Fv /Fm ) indicated that both strains had an equal ability to process captured photons and deliver them to PSII reaction centers. However, CS506 grew faster than CS509. This was consistent with its higher light requirement for saturation of photosynthesis (Ik ). Greater shade tolerance of CS509 was indicated by its higher ability to harvest light (α), lower photosynthetic light compensation point (Ic ), and higher chlorophyll a to biovolume ratio. Strain-specific differences were found in relation to non-photochemical quenching, effective absorption cross-sectional area of PSIIα-centers (σPSIIα), and the antenna connectivity parameter of PSIIα (Jcon PSIIα). These findings highlighted differences in the transfer of excitation from phycobilisome/PSII to PSI, on the dependence on different pigments for light harvesting and on the functioning of the PSII reaction centers between the two strains. The results of this study showed that both performance and composition of the photosynthetic apparatus are different between these strains, though with only two strains examined we cannot attribute the performance of strain 506 to its ability to produce cylindrospermopsins. The emphasis on a strain-specific light adaptation/acclimation is crucial to our understanding of how different light conditions (both quantity and quality) can trigger the occurrence of different C. raciborskii strains and control their competition and/or dominance in natural ecosystems. © 2013 Phycological Society of America.
Construction costs, chemical composition and payback time of high- and low-irradiance leaves
The effect of irradiance on leaf construction costs, chemical composition, and on the payback time of leaves was investigated. To enable more generalized conclusions, three different systems were studied: top and the most-shaded leaves of 10 adult tree species in a European mixed forest, top leaves of sub-dominant trees of two evergreen species growing in small gaps or below the canopy in an Amazonian rainforest, and plants of six herbaceous and four woody species grown hydroponically at low or high irradiance in growth cabinets. Daily photon irradiance varied 3-6-fold between low- and high-light leaves. Specific leaf area (SLA) was 30-130% higher at low light. Construction costs, on the other hand, were 1-5% lower for low-irradiance leaves, mainly because low-irradiance leaves had lower concentrations of soluble phenolics. Photosynthetic capacity and respiration, expressed per unit leaf mass, were hardly different for the low- and high-light leaves. Estimates of payback times of the high-irradiance leaves ranged from 2-4 d in the growth cabinets, to 15-20 d for the adult tree species in the European forest. Low-irradiance leaves had payback times that were 2-3 times larger, ranging from 4 d in the growth cabinets to 20-80 d at the most shaded part of the canopy of the mixed forest. In all cases, estimated payback times were less than half the life span of the leaves, suggesting that even at time-integrated irradiances lower than 5% of the total seasonal value, investment in leaves is still fruitful from a carbon-economy point of view. A sensitivity analysis showed that increased SLA of low-irradiance leaves was the main factor constraining payback times. Acclimation in the other five factors determining payback time, namely construction costs, photosynthetic capacity per unit leaf mass, respiration per unit leaf mass, apparent quantum yield, and curvature of the photosynthetic light-response-curve, were unimportant when the observed variation in each factor was examined.
Leaf attributes in the seasonally dry tropics a comparison of four habitats in northern Australia
DOI:10.1046/j.1365-2435.2003.00761.x
URL
[本文引用: 1]
\n\nPatterns of leaf attributes were compared at regional and global scales in relation to the seasonal availability of water.
From tropics to tundra: global convergence in plant functioning
Despite striking differences in climate, soils, and evolutionary history among diverse biomes ranging from tropical and temperate forests to alpine tundra and desert, we found similar interspecific relationships among leaf structure and function and plant growth in all biomes. Our results thus demonstrate convergent evolution and global generality in plant functioning, despite the enormous diversity of plant species and biomes. For 280 plant species from two global data sets, we found that potential carbon gain (photosynthesis) and carbon loss (respiration) increase in similar proportion with decreasing leaf life-span, increasing leaf nitrogen concentration, and increasing leaf surface area-to-mass ratio. Productivity of individual plants and of leaves in vegetation canopies also changes in constant proportion to leaf life-span and surface area-to-mass ratio. These global plant functional relationships have significant implications for global scale modeling of vegetation-atmosphere CO2 exchange.
Congruency analysis of species ranking based on leaf traits: Which traits are the more reliable?
DOI:10.1023/B:VEGE.0000046056.94523.57 URL [本文引用: 1]
Leaf economic traits from fossils support a weedy habit for early angiosperms
DOI:10.3732/ajb.0900290
PMID:21622407
[本文引用: 1]
Many key aspects of early angiosperms are poorly known, including their ecophysiology and associated habitats. Evidence for fast-growing, weedy angiosperms comes from the Early Cretaceous Potomac Group, where angiosperm fossils, some of them putative herbs, are found in riparian depositional settings. However, inferences of growth rate from sedimentology and growth habit are somewhat indirect; also, the geographic extent of a weedy habit in early angiosperms is poorly constrained. Using a power law between petiole width and leaf mass, we estimated the leaf mass per area (LMA) of species from three Albian (110-105 Ma) fossil floras from North America (Winthrop Formation, Patapsco Formation of the Potomac Group, and the Aspen Shale). All LMAs for angiosperm species are low (<125 g/m(2); mean = 76 g/m(2)) but are high for gymnosperm species (>240 g/m(2); mean = 291 g/m(2)). On the basis of extant relationships between LMA and other leaf economic traits such as photosynthetic rate and leaf lifespan, we conclude that these Early Cretaceous landscapes were populated with weedy angiosperms with short-lived leaves (<12 mo). The unrivalled capacity for fast growth observed today in many angiosperms was in place by no later than the Albian and likely played an important role in their subsequent ecological success.
Nutrient cycling and plant-soil feedbacks along a precipitation gradient in lowland Panama
DOI:10.1017/S0266467405002464
URL
[本文引用: 1]
This study addresses patterns of nutrient dynamics on a precipitation gradient (1800–3500 mm y−1) in lowland tropical forest with heterogeneous soil parent material, high plant species diversity and large changes in species composition. Mean foliar concentrations of phosphorus, potassium, calcium and magnesium decreased with increasing precipitation, whereas foliar carbon:nitrogen increased with increasing precipitation. Mean foliar nitrogen:phosphorus varied from 16.4–23.8 suggesting that plant productivity at these sites is limited by phosphorus. Total soil nitrogen increased as a function of foliar litter lignin:nitrogen, whereas net nitrogen mineralization rates decreased with increasing lignin:N indicating that as litter quality decreases, more soil nitrogen is held in soil organic matter and the mineralization of that nitrogen is slower. Extractable phosphorus in soil was negatively correlated with foliar litter lignin:phosphorus, illustrating effects of litter quality on soil phosphorus availability. Overall, the results suggest that variation in plant community composition along this precipitation gradient is tightly coupled with soil nutrient cycling. Much of our understanding of effects of precipitation on nutrient cycling in tropical forest is based on precipitation gradients across montane forest in Hawaii, where species composition and soil parent material are constant. Our results suggest that variation in parent material or species composition may confound predictions developed in model island systems.
Decline of leaf hydraulic conductance with dehydration: relationship to leaf size and venation architecture
DOI:10.1104/pp.111.173856
PMID:21511989
[本文引用: 1]
Across plant species, leaves vary enormously in their size and their venation architecture, of which one major function is to replace water lost to transpiration. The leaf hydraulic conductance (K(leaf)) represents the capacity of the transport system to deliver water, allowing stomata to remain open for photosynthesis. Previous studies showed that K(leaf) relates to vein density (vein length per area). Additionally, venation architecture determines the sensitivity of K(leaf) to damage; severing the midrib caused K(leaf) and gas exchange to decline, with lesser impacts in leaves with higher major vein density that provided more numerous water flow pathways around the damaged vein. Because xylem embolism during dehydration also reduces K(leaf), we hypothesized that higher major vein density would also reduce hydraulic vulnerability. Smaller leaves, which generally have higher major vein density, would thus have lower hydraulic vulnerability. Tests using simulations with a spatially explicit model confirmed that smaller leaves with higher major vein density were more tolerant of major vein embolism. Additionally, for 10 species ranging strongly in drought tolerance, hydraulic vulnerability, determined as the leaf water potential at 50% and 80% loss of K(leaf), was lower with greater major vein density and smaller leaf size (|r| = 0.85-0.90; P < 0.01). These relationships were independent of other aspects of physiological and morphological drought tolerance. These findings point to a new functional role of venation architecture and small leaf size in drought tolerance, potentially contributing to well-known biogeographic trends in leaf size.
Vegetation-mediated changes in microclimate reduce soil respiration as woodlands expand into grasslands
DOI:10.1890/03-0576 URL [本文引用: 1]
Leaf functional traits of ferns from different habitats in monsoon evergreen broad-leaved forest in Dinghushan Mountain
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
Vertical variation of leaf functional traits in temperate forest canopies in China
温带针阔混交林叶片性状随树冠垂直高度的变化规律
The photosynthetic capacity in 35 ferns and fern allies: mesophyll CO2 diffusion as a key trait
DOI:10.1111/nph.13719
PMID:26508678
[本文引用: 1]
Ferns and fern allies have low photosynthetic rates compared with seed plants. Their photosynthesis is thought to be limited principally by physical CO2 diffusion from the atmosphere to chloroplasts. The aim of this study was to understand the reasons for low photosynthesis in species of ferns and fern allies (Lycopodiopsida and Polypodiopsida). We performed a comprehensive assessment of the foliar gas-exchange and mesophyll structural traits involved in photosynthetic function for 35 species of ferns and fern allies. Additionally, the leaf economics spectrum (the interrelationships between photosynthetic capacity and leaf/frond traits such as leaf dry mass per unit area or nitrogen content) was tested. Low mesophyll conductance to CO2 was the main cause for low photosynthesis in ferns and fern allies, which, in turn, was associated with thick cell walls and reduced chloroplast distribution towards intercellular mesophyll air spaces. Generally, the leaf economics spectrum in ferns follows a trend similar to that in seed plants. Nevertheless, ferns and allies had less nitrogen per unit DW than seed plants (i.e. the same slope but a different intercept) and lower photosynthesis rates per leaf mass area and per unit of nitrogen. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
Soil fertility and water availability effects on trait dispersion and phylogenetic relatedness of tropical terrestrial ferns
DOI:10.1007/s00442-022-05131-w
PMID:35179630
[本文引用: 1]
Analysis of plant functional traits and their phylogenetic relationships has shed light on the processes structuring the occurrence patterns of angiosperm taxa across environmental gradients. In montane tropical forests, angiosperms coexist with diverse communities of terrestrial ferns, with distinct evolutionary histories, leaf morphology, and reproductive systems. Here we examined the functional traits, functional dispersion, and phylogenetic diversity of ferns across a well-described gradient of moisture and soil nutrient availability in a premontane tropical rainforest in western Panama. We measured 15 functional traits from 33 terrestrial fern species occurring in 12 one-ha plots. We applied RLQ and fourth-corner analyses to assess relationships between trait and environmental variables and used beta regression to evaluate how functional dispersion responds to environmental factors. In addition, we analyzed trait distributions with respect to fern phylogeny. We found that functional composition was predicted by soil variables and dry season rainfall. Leaf phosphorus (P) increased and leaf carbon (C) to nitrogen (N) ratio decreased with increasing soil total N:P ratio. Functional dispersion decreased with increasing soil total N:P in wet sites and with increasing manganese in dry sites, suggesting that low soil fertility and dry season moisture stress both tend to reduce functional diversity. Traits exhibited phylogenetic clustering primarily at deep nodes associated with tree versus herbaceous fern clades. Our results indicate that environmental filtering of functional traits affects ferns in a similar way to angiosperms and highlight the association of the early tree fern clade with low fertility soils.© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.
Let the concept of trait be functional!
DOI:10.1111/oik.2007.116.issue-5 URL [本文引用: 1]
Contrasting leaf-trait strategies in dominant liana and tree species of Indian tropical dry evergreen forest
DOI:10.1016/j.flora.2018.11.002 URL [本文引用: 1]
How does moss photosynthesis relate to leaf and canopy structure? Trait relationships for 10 Hawaiian species of contrasting light habitats
DOI:10.1111/j.1469-8137.2009.03061.x
PMID:19863726
[本文引用: 1]
Mosses are an understudied group of plants that can potentially confirm or expand principles of plant function described for tracheophytes, from which they diverge strongly in structure. We quantified 35 physiological and morphological traits from cell-, leaf- and canopy-level, for 10 ground-, trunk- and branch-dwelling Hawaiian species. We hypothesized that trait values would reflect the distinctive growth form and slow growth of mosses, but also that trait correlations would be analogous to those of tracheophytes. The moss species had low leaf mass per area and low gas exchange rate. Unlike for tracheophytes, light-saturated photosynthetic rate per mass (A(mass)) did not correlate with habitat irradiance. Other photosynthetic parameters and structural traits were aligned with microhabitat irradiance, driving an inter-correlation of traits including leaf area, cell size, cell wall thickness, and canopy density. In addition, we found a coordination of traits linked with structural allocation, including costa size, canopy height and A(mass). Across species, A(mass) and nitrogen concentration correlated negatively with canopy mass per area, analogous to linkages found for the 'leaf economic spectrum', with canopy mass per area replacing leaf mass per area. Despite divergence of mosses and tracheophytes in leaf size and function, analogous trait coordination has arisen during ecological differentiation.
Effects of crown development on leaf irradiance, leaf morphology and photosynthetic capacity in a peach tree
The three-dimensional (3-D) architecture of a peach tree (Prunus persica L. Batsch) growing in an orchard near Avignon, France, was digitized in April 1999 and again four weeks later in May 1999 to quantify increases in leaf area and crown volume as shoots developed. A 3-D model of radiation transfer was used to determine effects of changes in leaf area density and canopy volume on the spatial distribution of absorbed quantum irradiance (PAR(a)). Effects of changes in PAR(a) on leaf morphological and physiological properties were determined. Leaf mass per unit area (M(a)) and leaf nitrogen concentration per unit leaf area (N(a)) were both nonlinearly related to PAR(a), and there was a weak linear relationship between leaf nitrogen concentration per unit leaf mass (N(m)) and PAR(a). Photosynthetic capacity, defined as maximal rates of ribulose-1,5-bisphosphate carboxylase (Rubisco) carboxylation (V(cmax)) and electron transport (J(max)), was measured on leaf samples representing sunlit and shaded micro-environments at the same time that the tree crown was digitized. Both V(cmax) and J(max) were linearly related to N(a) during May, but not in April when the range of N(a) was low. Photosynthetic capacity per unit N(a) appeared to decline between April and May. Variability in leaf nitrogen partitioning between Rubisco carboxylation and electron transport was small, and the partitioning coefficients were unrelated to N(a). Spatial variability in photosynthetic capacity resulted from acclimation to varying PAR(a) as the crown developed, and acclimation was driven principally by changes in M(a) rather than the amount or partitioning of leaf nitrogen.
Towards a universal model for carbon dioxide uptake by plants
DOI:10.1038/s41477-017-0006-8
PMID:29150690
[本文引用: 1]
Gross primary production (GPP)-the uptake of carbon dioxide (CO) by leaves, and its conversion to sugars by photosynthesis-is the basis for life on land. Earth System Models (ESMs) incorporating the interactions of land ecosystems and climate are used to predict the future of the terrestrial sink for anthropogenic CO. ESMs require accurate representation of GPP. However, current ESMs disagree on how GPP responds to environmental variations, suggesting a need for a more robust theoretical framework for modelling. Here, we focus on a key quantity for GPP, the ratio of leaf internal to external CO (χ). χ is tightly regulated and depends on environmental conditions, but is represented empirically and incompletely in today's models. We show that a simple evolutionary optimality hypothesis predicts specific quantitative dependencies of χ on temperature, vapour pressure deficit and elevation; and that these same dependencies emerge from an independent analysis of empirical χ values, derived from a worldwide dataset of >3,500 leaf stable carbon isotope measurements. A single global equation embodying these relationships then unifies the empirical light-use efficiency model with the standard model of C photosynthesis, and successfully predicts GPP measured at eddy-covariance flux sites. This success is notable given the equation's simplicity and broad applicability across biomes and plant functional types. It provides a theoretical underpinning for the analysis of plant functional coordination across species and emergent properties of ecosystems, and a potential basis for the reformulation of the controls of GPP in next-generation ESMs.
Differences in leaf phenological traits between trees and shrubs are closely related to functional traits in a temperate forest
Nutrient resorption of two evergreen shrubs in response to long-term fertilization in a bog.
DOI:10.1007/s00442-013-2784-7 URL [本文引用: 1]
Intraspecific trait variation in plants: a renewed focus on its role in ecological processes
DOI:10.1093/aob/mcab011
PMID:33507251
[本文引用: 1]
Investigating the causes and consequences of intraspecific trait variation (ITV) in plants is not novel, as it has long been recognized that such variation shapes biotic and abiotic interactions. While evolutionary and population biology have extensively investigated ITV, only in the last 10 years has interest in ITV surged within community and comparative ecology.Despite this recent interest, still lacking are thorough descriptions of ITV's extent, the spatial and temporal structure of ITV, and stronger connections between ITV and community and ecosystem properties. Our primary aim in this review is to synthesize the recent literature and ask: (1) How extensive is intraspecific variation in traits across scales, and what underlying mechanisms drive this variation? (2) How does this variation impact higher-order ecological processes (e.g. population dynamics, community assembly, invasion, ecosystem productivity)? (3) What are the consequences of ignoring ITV and how can these be mitigated? and (4) What are the most pressing research questions, and how can current practices be modified to suit our research needs? Our secondary aim is to target diverse and underrepresented traits and plant organs, including anatomy, wood, roots, hydraulics, reproduction and secondary chemistry. In addressing these aims, we showcase papers from the Special Issue.Plant ITV plays a key role in determining individual and population performance, species interactions, community structure and assembly, and ecosystem properties. Its extent varies widely across species, traits and environments, and it remains difficult to develop a predictive model for ITV that is broadly applicable. Systematically characterizing the sources (e.g. ontogeny, population differences) of ITV will be a vital step forward towards identifying generalities and the underlying mechanisms that shape ITV. While the use of species means to link traits to higher-order processes may be appropriate in many cases, such approaches can obscure potentially meaningful variation. We urge the reporting of individual replicates and population means in online data repositories, a greater consideration of the mechanisms that enhance and constrain ITV's extent, and studies that span sub-disciplines.© The Author(s) 2021. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.
Understanding ecological variation across species area-based vs mass- based expression of leaf traits
DOI:10.1111/nph.12345 PMID:23692294 [本文引用: 1]
Specific leaf area and leaf dry matter content as alternative predictors of plant strategies
DOI:10.1046/j.1469-8137.1999.00427.x
URL
[本文引用: 1]
A key element of most recently proposed plant strategy schemes is an axis of resource capture, usage and availability. In the search for a simple, robust plant trait (or traits) that will allow plants to be located on this axis, specific leaf area is one of the leading contenders. Using a large new unpublished database, we examine the variability of specific leaf area and other leaf traits, the relationships between them, and their ability to predict position on the resource use axis. Specific leaf area is found to suffer from a number of drawbacks; it is both very variable between replicates and much influenced by leaf thickness. Leaf dry‐matter content (sometimes referred to as tissue density) is much less variable, largely independent of leaf thickness and a better predictor of location on an axis of resource capture, usage and availability. However, it is not clear how useful dry matter content will be outside northwest Europe, and in particular in dry climates with many succulents.
The worldwide leaf economics spectrum
DOI:10.1038/nature02403 [本文引用: 2]
Vertical variation and economic strategy of leaf trait of major tree species in a typical mixed broadleaved-Korean pine forest
DOI:10.17521/cjpe.2019.0307 URL [本文引用: 2]
典型阔叶红松林主要树种叶性状的垂直变异及经济策略
Quantifying leaf-trait covariation and its controls across climates and biomes
DOI:10.1111/nph.2019.221.issue-1 URL [本文引用: 2]
Effects of forest patch type and site on herb-layer vegetation in a temperate forest ecosystem
DOI:10.1016/j.foreco.2012.12.039 URL [本文引用: 1]
Effects of groundwater salinities on photosynthesis and water consumption characteristics of Tamarix chinensis in the Yellow River Delta
DOI:10.31497/zrzyxb.20191209 URL [本文引用: 1]
地下水矿化度对黄河三角洲柽柳光合及耗水特征的影响
DOI:10.31497/zrzyxb.20191209
[本文引用: 1]
为了解黄河三角洲地下水浅埋区柽柳(Tamarix chinensis Lour.)对地下水矿化度的适应特征,运用叶片气体交换和树干液流技术,测定了地下水埋深为0.9 m时淡水、微咸水(3 g·L<sup>-1</sup>)、咸水(8 g·L<sup>-1</sup>)和盐水(20 g·L<sup>-1</sup>)四种矿化度下柽柳的光合作用和水分利用等参数。结果表明:随地下水矿化度升高:(1)土壤水、盐含量和土壤溶液绝对浓度均逐渐升高。(2)叶片最大净光合速率和光饱和点在咸水下最高;表观量子效率、气孔导度、蒸腾速率、胞间CO<sub>2</sub>浓度、树干液流速率等在微咸水下最高;以上指标均在盐水矿化度下最低。水分利用效率和气孔限制值在微咸水下最低,盐水下最高。在地下水埋深0.9 m时,咸水矿化度下柽柳光合效率高,光照生态幅宽,水分利用效率高,适于柽柳幼苗的生长。
Leaf functional traits of Ceratoides latens in northern slope of Kunlun Mountain and its regional difference with the altitude
昆仑山北坡驼绒藜叶片功能性状及其海拔差异性
以昆仑山北坡前山带驼绒藜为试验材料,测定其光合、叶片氮、磷含量及比叶面积,并检验其相关关系。研究结果表明:①光合有效辐射和土壤含水量随海拔升高而增大,温度随海拔升高而降低;②驼绒藜的光合日进程曲线为“单峰型”,且没有出现明显的“午休”现象,到上午10:00左右达到最大值,而后逐渐下降;其最大净光合速率随海拔升高而增大;③随着海拔上升,叶片氮、磷含量及比叶面积均呈升高趋势;④单位面积最大光合速率(Aarea)与叶片氮(Nmass)、磷(Pmass)含量及比叶面积(SLA)的相关性均达到极显著水平,且呈高度正相关。逐步回归分析发现对Aarea贡献大小顺序为Nmass>SLA>Pmass。
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