Chin J Plant Ecol ›› 2019, Vol. 43 ›› Issue (2): 131-138.doi: 10.17521/cjpe.2019.0291

• Research Articles • Previous Articles     Next Articles

Conduits anatomical structure and leaf traits of diffuse- and ring-porous stems in subtropical evergreen broad-leaved forests

ZHANG Zhen-Zhen1,*(),ZHAO Ping2,ZHANG Jin-Xiu1,SI Yao1   

  1. 1 School of Geography and Environmental Sciences, Zhejiang Normal University, Jinhua, Zhejiang 231004, China
    2 South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China
  • Received:2018-11-14 Accepted:2019-01-30 Online:2019-06-04 Published:2019-02-20
  • Contact: ZHANG Zhen-Zhen
  • Supported by:
    Supported by the National Natural Science Foundation of China(41630752);Supported by the National Natural Science Foundation of China(41701226);the Zhejiang Province Public Welfare Technology Application Research Project(GF19C030003)

Abstract: <i>Aims</i>

The conduits characters are critical for plants to develop their survival strategies. Our current knowledge in this regard remains limited for the subtropical forest. In this study, we set a study objective to quantify the relationship between the conduits characters and the leaf functional traits of the dominant species in the region.


Two dominant species, Castanopsis chinensis and Schima superba, in a subtropical forest in Shimentai Nature Reserve were selected to compare their differences in functional traits, including conduits anatomical structure, the leaf morphological characteristics, and leaf physiological characteristics. The study was conducted during the dry season (October to March of the following year) for quantifying the ring-porous and diffuse-porous species. A series of t-tests were performed to quantify the statistical differences of all traits between the two species.

<i>Important findings</i>

We found that the density of conduits of S. superba (diffuse-porous) was significantly higher than that of C. chinensis (ring-porous), while the diameter of conduits for C. chinensis was much larger than that of S. superba. The leaf water content and the Chlorophyll a/Chlorophyll b ratio were much higher for S. superba than that of C. chinensis; the stomatal density and specific leaf area (SLA) tended to be higher in C. chinensis. In addition, it appeared that the differences in leaf specific net photosynthetic rates and the leaf stomatal conductance were not significant between S. superba and C. chinensis. These results indicated that the ring-porous species C. chinensis maintain a high photosynthetic capacity by maintaining a higher SLA at the expense of low leaf water content in responding to the water stress. The diffuse-porous species S. superba, meanwhile, tended to maintain a high capability of light transform under drought. These functional differences might be responsible for the succession pathways under the gradual changes of global precipitation for the region.

Key words: ring-porous, diffuse-porous, leaf trait, survival strategy, subtropical forest

Table 1

Community characteristics of the two dominant tree species of the subtropical forest in Shimentai of Guangdong Province"


Tree height
(mean ± SE) (m)
Castanopsis chinensis
壳斗科 Fagaceae 0.22 0.06 0.04 0.32 137 13.2 ± 2.1
木荷 Schima superba 山茶科 Theaceae 0.15 0.03 0.02 0.20 69 15.4 ± 1.3

Fig. 1

Microscopic images of conduits in Schima superba and Castanopsis chinensis."

Fig. 3

Specific leaf area (SLA), leaf water content (LWC), stomatal quantity (Sd) and size (Ss) of Schima superba and Castanopsis chinensis (mean ± SE). *, p < 0.05; **, p < 0.01."

Fig. 4

Leaf chlorophyll (Chl) content, Chl a/b, net photosynthetic rate (Pn), stomatal conductance (Cond), leaf transpiration rate (Tr) and instantaneous water use efficiency (WUEi) for Schima superba and Castanopsis chinensis (mean ± SE). *, p < 0.05; **, p < 0.01."

Fig. 2

Mean (± SE) of conduit density and diameter for Schima superba and Castanopsis chinensis. **, p < 0.01."

[1] Arnon DI ( 1949). Copper enzymes in isolated chloroplasts. Polyphenoloxidase in Beta vulgaris. Plant Physiology, 24, 1-15.
doi: 10.1104/pp.24.1.1
[2] Blonder B, Vasseur F, Violle C, Shipley B, Enquist BJ, Vile D ( 2015). Testing models for the leaf economics spectrum with leaf and whole-plant traits in Arabidopsis thaliana. AoB Plants, 7, plv049. DOI: 10.1093/aobpla/plv049.
[3] Breshears DD, Myers OB, Meyer CW, Barnes FJ, Zou CB, Allen CD, McDowell NG, Pockman WT ( 2009). Tree die-off in response to global change-type drought: Mortality insights from a decade of plant water potential measurements. Frontiers in Ecology and the Environment, 7, 185-189.
doi: 10.1890/080016
[4] Brodribb TJ, Holbrook NM, Edwards EJ, Gutiérrez MV ( 2010). Relations between stomatal closure, leaf turgor and xylem vulnerability in eight tropical dry forest trees. Plant, Cell & Environment, 26, 443-450.
[5] Büssis D, von Groll U, Fisahn J, Altmann T ( 2006). Stomatal aperture can compensate altered stomatal density in Arabidopsis thaliana at growth light conditions. Functional Plant Biology, 33, 1037-1043.
[6] Chave J, Coomes D, Jansen S, Lewis SL, Swenson NG, Zanne AE ( 2009). Towards a worldwide wood economics spectrum. Ecology Letters, 12, 351-366.
doi: 10.1111/ele.2009.12.issue-4
[7] Díaz S, Cabido M ( 2001). Vive la différence: Plant functional diversity matters to ecosystem processes. Trends in Ecology & Evolution, 16, 646-655.
[8] Díaz S, Cabido M, Casanoves F ( 1998). Plant functional traits and environmental filters at a regional scale. Journal of Vegetation Science, 9, 113-122.
doi: 10.2307/3237229
[9] Drake PL, Froend RH, Franks PJ ( 2013). Smaller, faster stomata: Scaling of stomatal size, rate of response, and stomatal conductance. Journal of Experimental Botany, 64, 495-505.
doi: 10.1093/jxb/ers347
[10] Esteban R, García-Plazaola JI, Hernández A, Fernández-Marín B ( 2018). On the recalcitrant use of Arnon’s method for chlorophyll determination. New Phytologist, 217, 474-476.
doi: 10.1111/nph.14932
[11] Fichot R, Chamaillard S, Depardieu C, Le Thiec D, Cochard H, Barigah TS, Brignolas F ( 2010). Hydraulic efficiency and coordination with xylem resistance to cavitation, leaf function, and growth performance among eight unrelated Populus deltoides× Populus nigra hybrids. Journal of Experimental Botany, 62, 2093-2106.
[12] Franks PJ, Drake PL, Beerling DJ ( 2009). Plasticity in maximum stomatal conductance constrained by negative correlation between stomatal size and density: An analysis using Eucalyptus globulus. Plant, Cell & Environment, 32, 1737-1748.
[13] Hulot FD, Lacroix G, Leschermoutoué F, Loreau M ( 2000). Functional diversity governs ecosystem response to nutrient enrichment. Nature, 405, 340-344.
doi: 10.1038/35012591
[14] Keyvan S ( 2010). The effects of drought stress on yield, relative water content, proline, soluble carbohydrates and chlorophyll of bread wheat cultivars. Journal of Animal and Plant Sciences, 8, 1051-1060.
[15] Klein T ( 2014). The variability of stomatal sensitivity to leaf water potential across tree species indicates a continuum between isohydric and anisohydric behaviours. Functional Ecology, 28, 1313-1320.
doi: 10.1111/1365-2435.12289
[16] Li K ( 2011). Community Structure and Leaf Characteristics of a Secondary Broad Leaved Forest in Hilly Area of Central Hunan, China. Master degree dissertation, Central South University of Forestry and Technology, Changsha.
[ 李凯 ( 2011). 湘中丘陵区次生阔叶林群落结构及叶片特征研究. 硕士学位论文, 中南林业科技大学, 长沙.]
[17] Li MC, Zhu JJ, Sun YR ( 2009). Responses of specific leaf area of dominant tree species in Northeast China secondary forests to light intensity. Chinese Journal of Ecology , 28, 1437-1442.
[ 李明财, 朱教君, 孙一荣 ( 2009). 东北次生林主要树种比叶面积对光照强度的响应. 生态学杂志, 28, 1437-1442.]
[18] Li WJ, Zuo JQ, Song YL, Liu JP, Li Y, Shen YS, Li JX ( 2015). Changes in spatio temporal distribution of drought/flood disaster in southern China under global climate warming. Meteorological Monthly , 41, 261-271.
[ 李维京, 左金清, 宋艳玲, 刘景鹏, 李瑜, 沈雨旸, 李景鑫 ( 2015). 气候变暖背景下我国南方旱涝灾害时空格局变化. 气象, 41, 261-271.]
[19] Matheny AM, Bohrer G, Vogel CS, Morin TH, He L, Frasson RPDM, Mirfenderesgi G, Schäfer KVR, Gough CM, Ivanov VY ( 2014). Species-specific transpiration responses to intermediate disturbance in a northern hardwood forest. Journal of Geophysical Research Biogeosciences, 119, 2292-2311.
doi: 10.1002/jgrg.v119.12
[20] Milcu A, Allan E, Roscher C, Jenkins T, Meyer ST, Flynn D, Bessler H, Buscot F, Engels C, Gubsch M ( 2013). Functionally and phylogenetically diverse plant communities key to soil biota. Ecology, 94, 1878-1885.
doi: 10.1890/12-1936.1
[21] Niu CY, Meinzer FC, Hao GY ( 2017). Divergence in strategies for coping with winter embolism among co-occurring temperate tree species: The role of positive xylem pressure, wood type and tree stature. Functional Ecology, 31, 1550-1560.
doi: 10.1111/fec.2017.31.issue-8
[22] Pérezharguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bretharte MS, Cornwell WK, Craine JM, Gurvich DE ( 2013). New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany, 61, 167-234.
doi: 10.1071/BT12225
[23] Peters EB, Mcfadden JP, Montgomery RA ( 2015). Biological and environmental controls on tree transpiration in a suburban landscape. Journal of Geophysical Research Biogeosciences, 115, G04006. DOI: 10.1029/2009JG001266.
[24] Poorter L, Mcdonald I, Alarcón A, Fichtler E, Licona JC, Peña-Claros M, Sterck F, Villegas Z, Sass-Klaassen U ( 2010). The importance of wood traits and hydraulic conductance for the performance and life history strategies of 42 rainforest tree species. New Phytologist, 185, 481-492.
doi: 10.1111/j.1469-8137.2009.03092.x
[25] Sack L, Holbrook NM ( 2006). Leaf hydraulics. Annual Review of Plant Biology, 57, 361-381.
doi: 10.1146/annurev.arplant.56.032604.144141
[26] Sperry JS, Nichols KL, Sullivan JEM, Eastlack SE ( 1994). Xylem embolism in ring-porous, diffuse-porous, and coniferous trees of northern Utah and interior Alaska. Ecology, 75, 1736-1752.
doi: 10.2307/1939633
[27] Sperry JS, Pockman WT ( 1993). Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis. Plant, Cell & Environment, 16, 279-287.
[28] Steppe K, Lemeur R ( 2007). Effects of ring-porous and diffuse-‌porous stem wood anatomy on the hydraulic parameters used in a water flow and storage model. Tree Physiology, 27, 43-52.
doi: 10.1093/treephys/27.1.43
[29] Takahashi S, Okada N, Nobuchi T ( 2013). Relationship between the timing of vessel formation and leaf phenology in ten ring-porous and diffuse-porous deciduous tree species. Ecological Research, 28, 615-624.
doi: 10.1007/s11284-013-1053-x
[30] Tateishi M, Kumagai TO, Utsumi Y, Umebayashi T, Shiiba Y, Inoue K, Kaji K, Cho K, Otsuki K ( 2008). Spatial variations in xylem sap flux density in evergreen oak trees with radial-porous wood: Comparisons with anatomical observations. Trees, 22, 23-30.
doi: 10.1007/s00468-007-0165-8
[31] von Allmen EI, Sperry JS, Bush SE ( 2013). Contrasting whole- tree water use, hydraulics, and growth in a co-dominant diffuse-porous vs. ring-porous species pair. Trees, 29, 717-728.
[32] Wang J, Ives NE, Lechowicz MJ ( 1992). The relation of foliar phenology to xylem embolism in trees. Functional Ecology, 6, 469-475.
doi: 10.2307/2389285
[33] Wardle DA, Barker GM, Bonner KI, Nicholson KS ( 1998). Can comparative approaches based on plant ecophysiological traits predict the nature of biotic interactions and individual plant species effects in ecosystems? Journal of Ecology,
86, 405-420.
[34] Westoby M, Falster DS, Moles AT, And PAV, Wright IJ ( 2002). Plant ecological strategies: Some leading dimensions of variation between species. Annual Review of Ecology & Systematics, 33, 125-159.
[35] Wright IJ, Ackerly DD, Bongers F, Harms KE, Ibarra-‌Manriquez G, Martinez-Ramos M, Mazer SJ, Muller-‌Landau HC, Paz H, Pitman NCA, Poorter L, Silman MR, Vriesendorp CF, Webb CO, Westoby M, Wright SJ ( 2007). Relationships among ecologically important dimensions of plant trait variation in seven neotropical forests. Annals of Botany, 99, 1003-1015.
doi: 10.1093/aob/mcl066
[36] Yan DL, Ren YY, Lian JF, Sheng LJ ( 2012). Leaf functional traits and their associated characteristics of three dominant families. Journal of Forestry Engineering , 26(3), 34-37.
[ 闫道良, 任燕燕, 连俊方, 盛琳杰 ( 2012). 3个优势科树种叶功能性状及其关联特性. 林业工程学报, 26(3), 34-37.]
[37] Zeng XM, Zhao P, Ouyang L, Zhu LW, Ni GY, Zhao XH ( 2017). Soil water use and adaptive regulation of Schima superba in the dry and wet seasons. Journal of Tropical and Subtropical Botany, 25(2), 105-114.
[ 曾小敏, 赵平, 欧阳磊, 朱丽薇, 倪广艳, 赵秀华 ( 2017). 荷木对干湿季土壤水分的利用和适应性调节. 热带亚热带植物学报, 25(2), 105-114.]
[38] Zeng XL, Wang DW, Liu JP, Wang SS, Fan X ( 2015). Effects of slope orientation on apparent traits and chlorophyll content of three cold-season turfgrasses. Grass Science , 32, 1823-1831.
[ 曾晓琳, 王大伟, 刘金平, 王思思, 范宣 ( 2015). 坡向对3种冷季型草坪草表观性状及叶绿素含量的影响. 草业科学, 32, 1823-1831.]
[39] Zeng XP, Zhao P, Cai XA, Rao XQ, Liu H, Ma L, Li CH ( 2006). A preliminary study on the tolerance of 25 species of South Asian tropical plants. Journal of Beijing Forestry University , 28(4), 92-99.
[ 曾小平, 赵平, 蔡锡安, 饶兴权, 刘惠, 马玲, 李长洪 ( 2006). 25种南亚热带植物耐阴性的初步研究. 北京林业大学学报, 28(4), 92-99.]
[40] Zhang JL, Poorter L, Cao KF ( 2012). Productive leaf functional traits of Chinese savanna species. Plant Ecology, 213, 1449-1460.
doi: 10.1007/s11258-012-0103-8
[41] Zhou G, Wei X, Wu Y, Liu S, Huang Y, Yan J, Zhang D, Zhang Q, Liu J, Meng Z, Wang C, Chu G, Liu S, Tang X, Wang C ( 2011). Quantifying the hydrological responses to climate change in an intact forested small watershed in Southern China. Global Change Biology, 17, 3736-3746.
doi: 10.1111/j.1365-2486.2011.02499.x
[42] Zhu L, Hu Y, Zhao X, Zeng X, Zhao P, Zhang Z, Ju Y ( 2017). The impact of drought on sap flow of cooccurring Liquidambar formosana Hance and Quercus variabilis Blume in a temperate forest, Central China. Ecohydrology, 10, 1828. DOI: 10.1002/eco.1828.
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[1] Cui Gao;Yuxia Chen;Ying Bao;Min Feng;Anming Lu. Studies on Sexual Organs and Embryological Development Morphology of Speirantha gardenii (Convallariaceae)[J]. Chin Bull Bot, 2010, 45(06): 705 -712 .
[2] Jiang Gao-ming. The Impact of Globae Increasing of CO2 on Plants[J]. Chin Bull Bot, 1995, 12(04): 1 -7 .
[3] Zhang Jun Han Bi-wen. Advance in the Study of Histochemical Localization for[J]. Chin Bull Bot, 1995, 12(专辑3): 131 -142 .
[4] Tang Yan-cheng. A Short Guide to the International Code of Botanical Nomenclature V.[J]. Chin Bull Bot, 1984, 2(04): 51 -57 .
[5] Xu Ji. The Protective Protein of Nitrogenase Against Oxygen Damage-Fe-S Protein[J]. Chin Bull Bot, 1986, 4(12): 1 -4 .
[6] . [J]. Chin Bull Bot, 2001, 18(05): 633 .
[7] Huang Zhao-xiang;Zheng Zhen-gui and Zhu Du. Ecological Effect of Taxodium ascendens-Oryza sativa Ecosystem(I) The Growing Characteristic of Taxodium Ascendens in the Ecosystem[J]. Chin Bull Bot, 1996, 13(02): 48 -51 .
[8] GU Rui-Sheng;LIU Qun-Lu;CHEN Xue-Mei and JIANG Xiang-Ning. Comparison and Optimization of the Methods on Protein Extraction and SDS-PAGE in Woody Plants[J]. Chin Bull Bot, 1999, 16(02): 171 -177 .
[9] Jiang Gao-ming. LI-6400 Portable Photosynthesis System: Principle, Function, Basic Operation and Main Problems and Solutions During Measurement[J]. Chin Bull Bot, 1996, 13(增刊): 72 -76 .
[10] Li Ling;Luo Yun-xiu;He Jian-hui and Pan Rui-chi. Promoting the Formation of Adventitious Roots in Cutting of Some Woody Plants by GL Reagent[J]. Chin Bull Bot, 1996, 13(增刊): 63 -65 .