鲁东丘陵同质生境中11个树种叶解剖学特性比较
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摘要
叶片是植物应对环境变化较为敏感的器官,叶解剖结构在一定程度上也体现出了树木对特殊生境的适应能力.为阐明植物对干旱生境的适应特征,对鲁东丘陵石灰岩山地同质生境内11个树种叶片以及叶脉的解剖结构进行分析.结果显示,各树种均为异面型叶,该类型叶片在解剖结构上具有显著的栅栏组织和海绵组织区分.叶表皮厚度在不同树种之间无明显差异,但上表皮普遍较下表皮厚(上表皮平均厚度为34.66μm,下表皮平均厚度为17.87μm).分别按乔、灌、藤3种生活型进行统计,发现叶片上下表皮厚度在生活型之间差异并不显著.各树种叶片均具有较发达的栅栏组织,而海绵组织排列较为稀疏.栅栏组织厚度与海绵组织厚度平均比值为2.42.在叶脉解剖结构中,各树种1级叶脉木质部厚度与韧皮部比值在1.43-5.30之间,且在树种间有显著差异,但该比值在乔、灌和藤3种生活型之间的差异不显著.叶脉解剖性状与叶片解剖性状存在显著相关(P <0.01).栅栏组织厚度与海绵组织厚度比值、叶脉木质部厚度在一定程度上表征了树种对干旱生境的适应能力.本研究从叶解剖特性方面阐明了叶片对干旱生境的适应特征,可为未来干旱瘠薄山地植被修复树种选择提供参考.(图4表4参48附图2)
Leaves serve as important and sensitive organs to environment changes. Leaf anatomical traits reflect the adaptation of trees to some special environments. Eleven co-existing tree species from the common environment of the limestone mountain area in eastern Shandong Province were selected. The seven anatomical traits of leaves and leaf veins were examined and recorded, including the upper epidermis thickness, palisade tissue thickness, spongy tissue thickness, lower epidermis thickness, main vascular diameter, xylem thickness, and phloem thickness. The results showed that the leaves of the 11 tree species were heterofacial and the differentiation of palisade cells and spongy tissues was obvious in the leaf cross section.The leaf epidermis thickness did not show significant differences among tree species; however, the upper epidermis thickness was generally larger than the lower epidermis thickness. The average thickness of the upper epidermis of the 11 species was34.66 μm and the average thickness of the lower epidermis was 17.87 μm. The average thickness of leaf epidermis did not show obvious differences among arbor, shrubs, or vines. The obvious palisade tissues in all 11 tree species, although spongy tissues were arranged sparsely, could be considered an important index for the tolerance assessment of tree species to the arid environment. The average ratio of palisade tissue thickness to spongy tissue thickness of the 11 species was 2.42. Regarding the leaf vein anatomy, the ratios of xylem and phloem thickness in leaf veins were 1.43–5.30, showing significant differences among the 11 tree species. However, the ratios in three life type, arbor, shrub, and vine, did not differ. There were significant correlations among leaf vein diameter, xylem thickness, and phloem thickness in the vein(P < 0.01). To a certain extent, the ratios of palisade tissues and spongy tissue thickness, the xylem thickness in leaf vein might serve as important indices for the tolerance assessment of tree species to the arid environment. This study shows some relative characteristics of leaves to arid habitats according to leaf anatomical traits, which would provide some references for the selection of tree species for the vegetation restoration in arid limestone mountain areas in the future.
Leaves serve as important and sensitive organs to environment changes. Leaf anatomical traits reflect the adaptation of trees to some special environments. Eleven co-existing tree species from the common environment of the limestone mountain area in eastern Shandong Province were selected. The seven anatomical traits of leaves and leaf veins were examined and recorded, including the upper epidermis thickness, palisade tissue thickness, spongy tissue thickness, lower epidermis thickness, main vascular diameter, xylem thickness, and phloem thickness. The results showed that the leaves of the 11 tree species were heterofacial and the differentiation of palisade cells and spongy tissues was obvious in the leaf cross section.The leaf epidermis thickness did not show significant differences among tree species; however, the upper epidermis thickness was generally larger than the lower epidermis thickness. The average thickness of the upper epidermis of the 11 species was34.66 μm and the average thickness of the lower epidermis was 17.87 μm. The average thickness of leaf epidermis did not show obvious differences among arbor, shrubs, or vines. The obvious palisade tissues in all 11 tree species, although spongy tissues were arranged sparsely, could be considered an important index for the tolerance assessment of tree species to the arid environment. The average ratio of palisade tissue thickness to spongy tissue thickness of the 11 species was 2.42. Regarding the leaf vein anatomy, the ratios of xylem and phloem thickness in leaf veins were 1.43–5.30, showing significant differences among the 11 tree species. However, the ratios in three life type, arbor, shrub, and vine, did not differ. There were significant correlations among leaf vein diameter, xylem thickness, and phloem thickness in the vein(P < 0.01). To a certain extent, the ratios of palisade tissues and spongy tissue thickness, the xylem thickness in leaf vein might serve as important indices for the tolerance assessment of tree species to the arid environment. This study shows some relative characteristics of leaves to arid habitats according to leaf anatomical traits, which would provide some references for the selection of tree species for the vegetation restoration in arid limestone mountain areas in the future.
引文
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2季子敬,全先奎,王传宽.兴安落叶松针叶解剖结构变化及其光合能力对气候变化的适应性[J].生态学报,2013,33(21):6967-6974[Ji ZJ,Quan XK,Wang CK.Variations in leaf anatomy of Larix gmelinii reflect adaptation of its photosynthetic capacity to climate changes[J].Acta Ecol Sin,2013,33(21):6967-6974]
3 He NP,Liu CC,Tian M,Li ML,Yang H,Yu GR,Guo DL,Smith MD,Yu Q,Hou JH.Variation in leaf anatomical traits from tropical to cold‐temperate forests and linkage to ecosystem functions[J].Func Ecol,2018,32(1):10-19
4张金玲,李玉灵,庞梦丽,朱辰光,毕志乐.臭柏异形叶解剖结构及其抗旱性的比较[J].西北植物学报,2017,37(9):1756-1763[Zhang JL,Li YL,Pang ML,Zhu CG,Bi ZL.Comparison of drought resistance of anatomic structure in the heterophylly mechanism of Sabina vulgaris[J].Acta Bot Bor-occid Sin,2017,37(9):1756-1763]
5 Hughes J,Hepworth C,Dutton C,Dunn JA,Hunt L,Stephens J,Cameron D,Waugh R,Gray JE.Reducing stomatal density in barley improves drought tolerance without impacting on yield[J].Plant Physiol,2017,174(2):776-787
6王常顺,汪诗平.植物叶片性状对气候变化的响应研究进展[J].植物生态学报,2015,39(2):206-216[Wang CS,Wang SP.A review of research on responses of leaf traits to climate change[J].Chin J Plant Ecol,2015,39(2):206-216]
7 Kr?ber W,Heklau H,Bruelheide H.Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits[J].Plant Biol,2015,17(2):373-383
8 Niinemetsü,Portsmuth A,Tobias M.Leaf shape and venation pattern alter the support investments within leaf lamina in temperate species:a neglected source of leaf physiological differentiation?[J].Funct Ecol,2007,21(1):28-40
9 Sack L,Cowan PD,Holbrook NM.The major veins of mesomorphic leaves revisited:tests for conductive overload in Acer saccharum(Aceraceae)and Quercus rubra(Fagaceae)[J].Am J Bot,2003,90(1):32-39
10 Sack L,Scoffoni C,Mckown AD,Frole K,Rawls M.Developmentally based scaling of leaf venation architecture explains global ecological patterns[J].Nat Commun,2012,3:837
11 Sack L,Holbrook NM.Leaf hydraulics[J].Annu Rev Plant Biol,2006,57:361-381
12 Sack L,Scoffoni C.Leaf venation:structure,function,development,evolution,ecology and applications in the past,present and future[J].New Phytol,2013,198(4):983-1000
13 Wright IJ,Reich PB,Westoby M,Ackerly DD,Baruch Z,Bongers F,Cavender-Bares J,Chapin T,Cornelissen JHC,Diemer M,Flexas J,Garnier E,Groom PK,Gulias J,Hikosaka K,Lamont B B,Lee T,Lee W,Lusk C,Midgley JJ,Navas ML,Niinemets U,Oleksyn J,Osada N,Poorter H,Poot P,Prior L,Pyankov VI,Roumet C,Thomas SC,Tjoelker MG,Veneklaas EJ,Villar R.The worldwide leaf economics spectrum[J].Nature,2004,428(6985):821-827
14陈莹婷,许振柱.植物叶经济谱的研究进展[J].植物生态学报,2014,38(10):1135-1153[Chen YT,Xu ZZ.Review on research of leaf economics spectrum[J].Chin J Plant Ecol,2014,38(10):1135-1153]
15李周,赵雅洁,宋海燕,张静,陶建平,刘锦春.不同水分处理下喀斯特土层厚度异质性对两种草本叶片解剖结构和光合特性的影响[J].生态学报,2018,38(2):721-732[Li Z,Zhao YJ,Song HY,Zhang J,Tao JP,Liu JC.The effects of soil thickness heterogeneity on the anatomical structure and photosynthetic characteristics of two herbaceous leaves under different water treatments[J].Acta Ecol Sin,2018,38(2):721-732]
16 Caringella MA,Bongers FJ,Sack L.Leaf hydraulic conductance varies with vein anatomy across Arabidopsis thaliana wild-type and leaf vein mutants[J].Plant Cell Environ,2015,38(12):2735-2746
17 Maréchaux I,Bartlett MK,Sack L,Baraloto C,Engel J,Joetzjer E,Chave J.Drought tolerance as predicted by leaf water potential at turgor loss point varies strongly across species within an Amazonian forest[J].Func Ecol,2015,29(10):1268-1277
18 Li L,Ma ZQ,Niinemetsü,Guo DL.Three key sub-leaf modules and the diversity of leaf designs[J].Front Plant Sci,2017(8):1542
19 Li L,McCormack ML,Ma CE,Kong D,Zhang Q,Chen XY,Zeng H,Niinemetsü,Guo DL.Leaf economics and hydraulic traits are decoupled in five species-rich tropical-subtropical forests[J].Ecol Lett,2015,18(9):899-906
20邵水仙,董智,李红丽,许婷婷.不同造林模式对退化石灰岩山地土壤理化性质及水文效应的影响[J].水土保持学报,2015,29(1):263-267[Shao SX,Dong Z,Li HL,Xu TT.Influence of different afforestation models on soil physical and chemical properties and hydrological effect in degradation limestone mountain[J].J Res Soil Water Conserv,2015,29(1):263-267]
21李小倩,杨吉华,魏晓明.鲁中南石灰岩山地针阔混交林土壤理化性状及水文效应[J].水土保持学报,2016,30(1):208-211[Li XQ,Yang JH,Wei XM.Soil Physical and chemical properties and hydrological effects of coniferous and broadleaved mixed forest in limestone mountains[J].J Res Soil Water Conserv,2016,30(1):208-211]
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