Comparison of Two Models Estimating Water Use Efficiency Based on δ13C: A Case Study of Plant Species in Leidongping, Emei Mountains of China
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摘要:
目的 通过对基于稳定碳同位素值（δ13C）估算水分利用效率（iWUE）的2种模型（是否考虑叶肉导度，gm）之间差异的研究，有助于进一步理解gm对植物iWUE的影响。 方法 本研究选择峨眉山雷洞坪针阔混交林中不同生长型的植物，共采集了48个物种的117个样品，通过测定叶片δ13C，对比了早期估算iWUE模型（iWUEsim模型，将gm看作无穷大）与包含gm效应的iWUE模型（iWUEmes模型）之间的iWUE差异（iWUEdifference），以及这种差异随植物生长型变化的趋势。 结果 结果显示，iWUEsim的平均值（42.23±1.33 μmol·mol−1）显著高于iWUEsem的平均值（28.10±0.65 μmol·mol−1），高估比例3.64%～72.11%（平均49.58±1.30%）。不同生长型植物的iWUEsim均显著大于iWUEsem，其中草本植物高估比例3.64%～48.00%（平均29.20±5.16%），灌木高估比例13.68%～67.73%（平均48.44±1.83%），乔木高估比例32.04%～72.11%（平均54.08±1.47%）。此外，叶片功能性状也对iWUEdifference有重要影响，相比于叶片厚度、叶片干物质含量和单位质量叶片氮含量，单位面积叶片氮含量和比叶重是影响iWUEdifference变化的主要因素。 结论 将gm看作无穷大会造成对基于δ13C估算的iWUE的高估，未来估算植物iWUE时应考虑gm的影响。 Abstract:Objective The study of differences between two models estimating intrinsic water use efficiency (iWUE) based on stable carbon isotope values (δ13C), considering leaf mesophyll conductance (gm) or not, contributes to a better understanding of the impact of gm on plant iWUE. Method In this study, we used different plant growth forms in the coniferous broadleaved mixed forest at Leidongping of Emei Mountains. By measuring the δ13C of leaves, we compared the difference in the calculated iWUE (iWUEdifference) between the early used iWUE model (iWUEsim model, assuming gm as infinity) and the newly proposed iWUE model with gm effect (iWUEmes model), and to detect whether this difference varies with plant growth forms. Result The results showed that the average iWUEsim was 42.23±1.33 μmol·mol−1, which was significantly higher than the average of iWUEsem (28.10±0.65 μmol·mol−1), with an overestimation ratio of 3.64%～72.11% (average of 49.58±1.30%). All the iWUEsim of different growth forms of plants were significantly higher than iWUEsem. The overestimation ratio was 3.64%～48.00% (average of 29.20±5.16%) for herbs, 13.68%～67.73% (average of 48.44±1.83%) for shrubs, and 32.04%～72.11% (average of 54.08±1.47%) for trees. In addition, the leaf functional traits also had important effects on the iWUEdifference. The leaf nitrogen content per unit area, the leaf mass per area were the main influencing factors for the variation of iWUEdifference, compared with the effects of leaf thickness, leaf dry matter content, and leaf nitrogen content per unit mass. Conclusion The assumption of gm as infinity will lead to an overestimation of iWUE based on δ13C, and the effect of gm should be considered in the future estimation of plant iWUE. -
图 1 不同生长型植物叶片的iWUE特征
注：图中s表示物种数，n表示样品数量。图中不同小写字母表示iWUEsim和iWUEsem存在显著差异（P＜0.05）。图中不同大写字母表示不同生长型植物间iWUE（iWUEsim，iWUEsem，iWUEdifference）存在显著差异（P＜0.05）。The s represents the number of species and n represents the number of samples. Different lowercase letters in the figure indicate significant differences in iWUEsim and iWUEsem (P＜0.05). Different uppercase letters in the figure indicate significant differences in iWUE (iWUEsim, iWUEsem, iWUEdifference) among plants of different growth forms (P＜0.05).
Figure 1. The characteristics of iWUE in leaf of different plant growth forms
图 2 不同生长型植物叶片的功能性状
注：LT，叶厚；LDMC，叶干物质含量；LMA，比叶重；Narea，单位面积叶片氮含量；Nmass，单位质量叶片氮含量。图中不同大写字母表示不同生长型植物间叶片功能性状差异显著（P＜0.05）。LT, leaf thickness; LDMC, leaf dry matter content; LMA, leaf mass per area; Narea, leaf nitrogen content per unit area; Nmass, leaf nitrogen content per unit mass. Different capital letters in the figure indicate significant differences in leaf functional traits between different growth types of plants (P＜0.05).
Figure 2. The leaf functional traits in different plant growth forms
图 3 叶片功能性状与iWUEdifference的关系
注：LT，叶厚；LDMC，叶干物质含量；LMA，比叶重；Narea，单位面积叶片氮含量；Nmass，单位质量叶片氮含量。LT, leaf thickness; LDMC, leaf dry matter content; LMA, leaf mass per area; Narea, leaf nitrogen content per unit area; Nmass, leaf nitrogen content per unit mass.
Figure 3. The relationship between leaf functional traits and iWUEdifference
图 4 叶片功能性状对iWUEdifference的相对重要性
注：LT，叶厚；LDMC，叶干物质含量；LMA，比叶重；Narea，单位面积叶片氮含量；Nmass，单位质量叶片氮含量。LT, leaf thickness; LDMC, leaf dry matter content; LMA, leaf mass per area; Narea, leaf nitrogen content per unit area; Nmass, leaf nitrogen content per unit mass.
Figure 4. The relative importance of leaf functional traits to iWUEdifference
表 1 峨眉山雷洞坪针阔混交林中所采集的不同生长型植物名录
Table 1. Catalogue of different growth forms of plants collections in coniferous broadleaved mixed forest in Leidongping, Mount Emei
植物生长型
Plant growth forms采集植物名称
Species names草本植物
Herbs黄水枝（Tiarella polyphylla） 、尖叶堇菜（Viola acutifolia） 、山酢浆草（Oxalis griffithii） 、酸模叶蓼（Persicaria lapathifolia） 、腋花扭柄花（Streptopus simplex） 灌木
Shrubs宝兴悬钩子（Rubus ourosepalus） 、峨眉蔷薇（Rosa omeiensis） 、峨眉卫矛（Euonymus omeiensis） 、枸骨（Ilex cornuta） 、合轴荚蒾（Viburnum sympodiale） 、寒莓（Rubus buergeri） 、华西忍冬（Lonicera webbiana） 、栎叶杜鹃（Rhododendron phaeochrysum） 、荚蒾（Viburnum dilatatum） 、冷地卫矛（Euonymus frigidus） 、南蛇藤（Celastrus orbiculatus） 、绒毛杜鹃（Rhododendron pachytrichum） 、陕甘花楸（Sorbus koehneana） 、山梅花（Philadelphus incanus） 、石枣子（Euonymus sanguineus） 、弯梗菝葜（Smilax aberrans） 、微毛樱桃（Prunus clarofolia） 、五味子（Schisandra chinensis） 、腺果杜鹃（Rhododendron davidii） 乔木
Trees糙皮桦（Betula utilis） 、长尾槭（Acer caudatum） 、川榛（Corylus heterophylla var. sutchuanensis） 、刺榛（Corylus ferox） 、灯笼树（Enkianthus chinensis） 、杜仲（Eucommia ulmoides） 、峨眉冷杉（Abies fabri） 、葛罗槭（Acer davidii） 、华西臭樱（Prunus hypoxantha） 、冷杉属（Abies） 、泡花树（Meliosma cuneifolia） 、青榨槭（Acer davidii） 、山鸡椒（Litsea cubeba） 、深裂中华槭（Acer sinense） 、石灰花楸（Sorbus folgneri） 、四川蜡瓣花（Corylopsis willmottiae） 、四蕊槭（Acer stachyophyllum） 、五裂槭（Acer oliverianum） 、五尖槭（Acer maximowiczii） 、细齿稠李（Prunus obtusata） 、银叶杜鹃（Rhododendron argyrophyllum） 、榆树（Ulmus pumila） 、紫花槭（Acer pseudosieboldianum） 、紫弹树（Celtis biondii） 
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