琵琶柴和沙拐枣茎的木质部结构的差异性及空间变异特征

doi:10.13866/j.azr.2023.12.12

干旱区研究 ›› 2023, Vol. 40 ›› Issue (12): 1996-2006.doi: 10.13866/j.azr.2023.12.12

琵琶柴和沙拐枣茎的木质部结构的差异性及空间变异特征

沈辉^1,²(),张静^1,³,彭兰^1,⁴,陶冶^1,³,臧永新^1,³,张元明^1,³()

1.中国科学院新疆生态与地理研究所荒漠与绿洲生态国家重点实验室，干旱区生态安全与可持续发展重点实验室，新疆乌鲁木齐 830011
2.中国科学院大学，北京 100093
3.中国科学院新疆生态与地理研究所新疆植物基因资源保护与利用重点实验室，新疆乌鲁木齐 830011
4.新疆大学生态与环境学院，新疆乌鲁木齐 830011

收稿日期:2023-04-06 修回日期:2023-06-21 出版日期:2023-12-15 发布日期:2023-12-18
通讯作者: 张元明. E-mail: zhangym@ms.xjb.ac.cn
作者简介:沈辉(1997-)，女，硕士研究生，主要从事荒漠植物生理生态研究. E-mail: shenhui20@mails.ucas.ac.an
基金资助:
中国科学院“西部之光-西部青年学者”项目(2021-XBQNXZ-006);国家自然科学基金联合重点基金(U2003214);新疆维吾尔自治区自然科学基金重点项目(2022D01D083)

Differences and spatial variation in the stem xylem structural traits of Reaumuria soongarica and Calligonum mongolicum

SHEN Hui^1,²(),ZHANG Jing^1,³,PENG Lan^1,⁴,TAO Ye^1,³,ZANG Yongxin^1,³,ZHANG Yuanming^1,³()

1. State Key Laboratory of Desert and Oasis Ecology, Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, Xinjiang, China
2. University of Chinese Academy of Science, Beijing 100093, China
3. Xinjiang Key Laboratory of Conservation and Utilization of Plant Gene Resources, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, Xinjiang, China
4. College of Ecology and Environment, Xinjiang University, Urumqi 830011, Xinjiang, China

Received:2023-04-06 Revised:2023-06-21 Online:2023-12-15 Published:2023-12-18

摘要/Abstract

摘要：

木质部结构性状是植物适应不同水分条件的基础，也是植物响应环境条件变化的核心属性之一。以西北荒漠区的典型灌木琵琶柴(Reaumuria soongarica)和沙拐枣(Calligonum mongolicum)为研究对象，对其木质部结构和水力功能性状进行对比分析，以期了解同一生境中不同物种的差异性或相似性，以及同一物种在不同环境条件下的木质部结构性状的可塑性。结果表明：（1）琵琶柴和沙拐枣的木质部结构性状存在显著差异，沙拐枣平均导管直径、脆弱性指数显著高于琵琶柴，而导管密度、导管分组指数则相反。（2）琵琶柴和沙拐枣的木质部结构性状随气候变化的规律具有差异，随年均降水量和干旱指数的增加，沙拐枣的平均导管直径与导管厚跨比显著下降，而琵琶柴的平均导管直径与导管厚跨比与之无关，仅理论导水率与年均降水量和干旱指数呈显著正相关关系。（3）琵琶柴木质部导水系统存在效率与安全的权衡，而沙拐枣中没有表现出效率和安全的权衡关系。（4）性状网络分析结果表明，琵琶柴和沙拐枣的中心性状均为平均导管直径，平均导管直径的变化介导了性状网络的变化。琵琶柴和沙拐枣的木质部结构性状存在显著差异，相对于沙拐枣，琵琶柴水分利用策略更加保守。

关键词: 优势灌木, 琵琶柴, 沙拐枣, 木质部结构, 植物性状网络, 气候因素, 生态适应策略

Abstract:

Xylem structure traits are the basis for plant adaptation to different water conditions and are core plant traits in response to changes in environmental conditions. The typical shrubs Reaumuria soongarica and Calligonum mongolicum in the northwest desert region were selected as the subjects for comparison between stem xylem structure and functional traits to understand the differences or similarities of different species in the same habitat and the xylem structure plasticity of the same species under different environmental conditions. The results showed that (1) Xylem structure traits of R. soongarica and C. mongolicum were significantly different. The mean vessel diameter and vulnerability index of C. mongolicum were significantly higher than those of R. soongarica, whereas the opposite was true for vessel density and vessel grouping index. (2) The patterns of xylem structure traits in response to climate change differed between R. soongarica and C. mongolicum. The mean vessel diameter and vessel thickness-to-span ratio of C. mongolicum significantly reduced with increasing mean annual precipitation and aridity index, whereas that of R. soongarica were unrelated, and only the theoretical hydraulic conductivity had a significant positive correlation with the mean annual precipitation and aridity index. (3) There was a trade-off between efficiency and safety in the xylem hydraulic conductivity system of R. soongarica, whereas none was observed in C. mongolicum. (4) The trait network analysis results indicated that the central traits of both R. soongarica and C. mongolicum were mean vessel diameters. Changes in mean vessel diameter mediate changes in the trait network. Xylem structure traits between R. soongarica and C. mongolicum were significantly different, with R. soongarica having a more conservative water use strategy than C. mongolicum.

Key words: tapical shrub, Reaumuria soongarica, Calligonum mongolicum, xylem structure, plant trait networks, climatic factors, ecological adaptation strategy

沈辉, 张静, 彭兰, 陶冶, 臧永新, 张元明. 琵琶柴和沙拐枣茎的木质部结构的差异性及空间变异特征[J]. 干旱区研究, 2023, 40(12): 1996-2006.

SHEN Hui, ZHANG Jing, PENG Lan, TAO Ye, ZANG Yongxin, ZHANG Yuanming. Differences and spatial variation in the stem xylem structural traits of Reaumuria soongarica and Calligonum mongolicum[J]. Arid Zone Research, 2023, 40(12): 1996-2006.

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参考文献 42

[1]	Pratt R B, Jacobsen A L. Conflicting demands on angiosperm xylem: Tradeoffs among storage, transport and biomechanics[J]. Plant, Cell & Environment, 2017, 40(6): 897-913.
[2]	Sperry J S, Love D M. What plant hydraulics can tell us about responses to climate-change droughts[J]. New Phytologist, 2015, 207(1): 14-27. doi: 10.1111/nph.13354 pmid: 25773898
[3]	Hacke U G, Jacobsen A L, Pratt R B. Vessel diameter and vulnerability to drought-induced embolism: Within-tissue and across-species patterns and the issue of survivorship bias[J]. Iawa Journal, 2022, 0(0): 1-16.
[4]	Jacobsen A L, Pratt R B, Venturas M D, et al. Large volume vessels are vulnerable to water-stress-induced embolism in stems of poplar[J]. Iawa Journal, 2019, 40(1): 4-22. doi: 10.1163/22941932-40190233
[5]	He P, Gleason S M, Wright I J, et al. Growing-season temperature and precipitation are independent drivers of global variation in xylem hydraulic conductivity[J]. Global Change Biology, 2020, 26(3): 1833-1841 doi: 10.1111/gcb.14929 pmid: 31749261
[6]	Olson M E, Soriano D, Rosell J A, et al. Plant height and hydraulic vulnerability to drought and cold[J]. Proceedings of the National Academy of Sciences, 2018, 115(29): 7551-7556. doi: 10.1073/pnas.1721728115
[7]	Jacobsen A L, Pratt R B, Davis S D, et al. Cavitation resistance and seasonal hydraulics differ among three arid Californian plant communities[J]. Plant, Cell & Environment, 2007, 30(12): 1599-1609.
[8]	Baas P, Carlquist S. A comparison of the ecological wood anatomy of the floras of Southern California and Israel[J]. Iawa Journal, 1985, 6(4): 349-353. doi: 10.1163/22941932-90000961
[9]	Zhang K, Yang D, Zhang Y, et al. Differentiation in stem and leaf traits among sympatric lianas, scandent shrubs and trees in a subalpine cold temperate forest[J]. Tree physiology, 2021, 41(11): 1992-2003. doi: 10.1093/treephys/tpab049
[10]	Jacobsen A L, Pratt R B, Tobin M F, et al. A global analysis of xylem vessel length in woody plants[J]. American Journal of Botany, 2012, 99(10): 1583-1591. doi: 10.3732/ajb.1200140 pmid: 22965850
[11]	Yao G, Nie Z, Turner N C, et al. Combined high leaf hydraulic safety and efficiency provides drought tolerance in Caragana species adapted to low mean annual precipitation[J]. New Phytologist, 2021, 229(1): 230-244. doi: 10.1111/nph.v229.1
[12]	Kocacinar F. Photosynthetic, hydraulic and biomass properties in closely related C₃ and C₄ species[J]. Physiologia Plantarum, 2015, 153(3): 454-466. doi: 10.1111/ppl.2015.153.issue-3
[13]	Morris H, Gillingham M A F, Plavcová L, et al. Vessel diameter is related to amount and spatial arrangement of axial parenchyma in woody angiosperms[J]. Plant, Cell & Environment, 2017, 41(1): 245-260.
[14]	Alonso-Forn D, Peguero-Pina J J, Ferrio J P, et al. Contrasting functional strategies following severe drought in two Mediterranean oaks with different leaf habit: Quercus faginea and Quercus ilex subsp. rotundifolia[J]. Tree physiology, 2021, 41(3): 371-387. doi: 10.1093/treephys/tpaa135 pmid: 33079165
[15]	黄海霞, 王刚, 陈年来. 荒漠灌木逆境适应性研究进展[J]. 中国沙漠, 2010, 30(5): 1060-1067.
	[Huang Haixia, Wang Gang, Chen Nianlai. Advance of studies on adaptation of desert shrub to environment stress[J]. Journal of Desert Research, 2010, 30(5): 1060-1067. ]
[16]	Shan L, Yang C, Li Y, et al. Effects of drought stress on root physiological traits and root biomass allocation of Reaumuria soongorica[J]. Acta Ecologica Sinica, 2015, 35(5): 155-159. doi: 10.1016/j.chnaes.2015.06.010
[17]	闫晴, 李菊艳, 尹忠东, 等. 典型株型沙生灌丛对风沙流场影响的数值模拟[J]. 干旱区研究, 2023, 40(5): 785-797.
	[Yan Qing, Li Juyan, Yin Zhongdong, et al. Numerical simulation of the influence of typical shrub types on wind-sand flow field[J]. Arid Zone Research, 2023, 40(5): 785-797. ]
[18]	Fan X, Yan X, Qian C, et al. Leaf size variations in a dominant desert shrub, Reaumuria soongarica, adapted to heterogeneous environments[J]. Ecology and Evolution, 2020, 10(18): 10076-10094. doi: 10.1002/ece3.v10.18
[19]	赵小仙, 李毅, 苏世平, 等. 3个地理种群蒙古沙拐枣同化枝解剖结构及抗旱性比较[J]. 中国沙漠, 2014, 34(5): 1293-1300. doi: 10.7522/j.issn.1000-694X.2013.00429
	[Zhao Xiaoxian, Li Yi, Su Shiping, et al. Drought resistance analysis based on anatomical structures of assimilating shoots of Calligonum mongolicum from three geographic populations[J]. Journal of Desert Research, 2014, 34(5): 1293-1300. ] doi: 10.7522/j.issn.1000-694X.2013.00429
[20]	Shen H, Zhang J, Peng L, et al. Spatial patterns and climatic factors influence the branch xylem anatomical traits of Reaumuria soongarica in the desert region of northwestern China[J]. Environmental and Experimental Botany, 2023, 210: 105338. doi: 10.1016/j.envexpbot.2023.105338
[21]	种培芳. 荒漠植物红砂、白刺和沙拐枣抗旱指标及抗旱性综合评价研究[D]. 兰州: 甘肃农业大学, 2010.
	[Zhong Peifang. Studies on Drought Resistant Indexs and Evaluated Drought Resistant Capability of Desert Plant Reaumuria soongarica, Nitraria tangutorum and Calligomum mongolicum[D]. Lanzhou: Gansu Agricultural University, 2010. ]
[22]	殷笑寒, 郝广友. 长白山阔叶树种木质部环孔和散孔结构特征的分化导致其水力学性状的显著差异[J]. 应用生态学报, 2018, 29(2): 352-360. doi: 10.13287/j.1001-9332.201802.035
	[Yin Xiaohan, Hao Guangyou. Divergence between ring-and diffuse-porous wood types in broadleaf trees of Changbai Mountains results in substantial differences in hydraulic traits[J]. Chinese Journal of Applied Ecology, 2018, 29(2): 352-360. ] doi: 10.13287/j.1001-9332.201802.035
[23]	Pfautsch S, Harbusch M, Wesolowski A, et al. Climate determines vascular traits in the ecologically diverse genus Eucalyptus[J]. Ecologe Letters, 2016, 19(3): 240-248.
[24]	Schuldt B, Knutzen F, Delzon S, et al. How adaptable is the hydraulic system of European beech in the face of climate change-related precipitation reduction?[J]. New Phytologist, 2016, 210(2): 443-458. doi: 10.1111/nph.13798 pmid: 26720626
[25]	刘润红, 白金连, 包含, 等. 桂林岩溶石山青冈群落主要木本植物功能性状变异与关联[J]. 植物生态学报, 2020, 44(8): 828-841. doi: 10.17521/cjpe.2019.0146
	[Liu Runhong, Bai Jinlian, Bao Han, et al. Variation and correlation in functional traits of main woody plants in the Cyclobalanopsis glauca community in the karst hills of Guilin, Southwest China[J]. Chinese Journal of Plant Ecology, 2020, 44(8): 828-841. ] doi: 10.17521/cjpe.2019.0146
[26]	Rosas T, Mencuccini M, Barba J, et al. Adjustments and coordination of hydraulic, leaf and stem traits along a water availability gradient[J]. New Phytologist, 2019, 223(2): 632-646. doi: 10.1111/nph.15684 pmid: 30636323
[27]	Bush S E, Pataki D E, Hultine K R, et al. Wood anatomy constrains stomatal responses to atmospheric vapor pressure deficit in irrigated, urban trees[J]. Oecologia, 2008, 156(1): 13-20. doi: 10.1007/s00442-008-0966-5 pmid: 18270747
[28]	Islam M, Rahman M, Bräuning A. Long-term hydraulic adjustment of three tropical moist forest tree species to changing climate[J]. Frontiers in Plant Science, 2018, 4(9):1-16.
[29]	García-Cervigón A I, Olano J M, von Arx G, et al. Xylem adjusts to maintain efficiency across a steep precipitation gradient in two coexisting generalist species[J]. Annals of Botany, 2018, 122(3): 461-472. doi: 10.1093/aob/mcy088 pmid: 29800073
[30]	Pivovaroff A L, Sack L, Santiago L S. Coordination of stem and leaf hydraulic conductance in southern California shrubs: A test of the hydraulic segmentation hypothesis[J]. New Phytologist, 2014, 203(3): 842-850. doi: 10.1111/nph.12850 pmid: 24860955
[31]	Meinzer F C, Mcculloh K A, Lachenbruch B, et al. The blind men and the elephant: The impact of context and scale in evaluating conflicts between plant hydraulic safety and efficiency[J]. Oecologia, 2010, 164(2): 287-296. doi: 10.1007/s00442-010-1734-x pmid: 20668883
[32]	Baas P, Ewers F W, Davis S D, et al. 15 - Evolution of Xylem Physiology[M]. Oxford: Academic Press, 2004: 273-295.
[33]	Liu H, Ye Q, Gleason S M, et al. Weak tradeoff between xylem hydraulic efficiency and safety: Climatic seasonality matters[J]. New Phytologist, 2021, 229(3): 1440-1452. doi: 10.1111/nph.16940 pmid: 33058227
[34]	Lens F, Sperry J S, Christman M A, et al. Testing hypotheses that link wood anatomy to cavitation resistance and hydraulic conductivity in the genus Acer[J]. New Phytologist, 2010, 190(3): 709-723. doi: 10.1111/nph.2011.190.issue-3
[35]	Hacke U G, Sperry J S, Pockman W T, et al. Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure[J]. Oecologia, 2001, 126(4): 457-461. doi: 10.1007/s004420100628 pmid: 28547229
[36]	方菁, 叶琳峰, 陈森, 等. 自然和人工生境被子植物枝木质部结构与功能差异[J]. 植物生态学报, 2021, 45(6): 650-658. doi: 10.17521/cjpe.2020.0430
	[Fang Jing, Ye Linfeng, Chen Sen, et al. Differences in anatomical structure and hydraulic function of xylem in branches of angiosperms in field and garden habitats[J]. Chinese Journal of Plant Ecology, 2021, 45(6): 650-658. ] doi: 10.17521/cjpe.2020.0430
[37]	刘玉冰, 张腾国, 李新荣, 等. 红砂(Reaumuria soongorica)忍耐极度干旱的保护机制:叶片脱落和茎中蔗糖累积[J]. 中国科学C辑: 生命科学, 2006, 36(4): 328-333.
	[Liu Yubing, Zhang Tengguo, Li Xinrong, et al. Reaumuria soongorica tolerates extreme drought protective mechanism: Leaf abscission and sucrose accumulation in stems[J]. Science in China Series C: Life Sciences, 2006, 36(4): 328-333. ]
[38]	Pandey S. Climatic influence on tree wood anatomy: A review[J]. Journal of Wood Science, 2021, 67(1): 24-31. doi: 10.1186/s10086-021-01956-w
[39]	Zhang S, Cao K, Fan Z, et al. Potential hydraulic efficiency in angiosperm trees increases with growth-site temperature but has no trade-off with mechanical strength[J]. Global Ecology and Biogeography, 2013, 22(8): 971-981. doi: 10.1111/geb.2013.22.issue-8
[40]	Gleason S M, Butler D W, Waryszak P. Shifts in leaf and stem hydraulic traits across aridity gradients in eastern Australia[J]. International Journal of Plant Sciences, 2013, 174(9): 1292-1301. doi: 10.1086/673239
[41]	彭丽萍, 戴岳, 师庆东. 新疆准东荒漠区5种典型植物水分来源[J]. 干旱区研究, 2018, 35(5): 1146-1152.
	[Peng Liping, Dai Yue, Shi Qingdong. Water sources of five typical plant species in desert area of the Eastern Junggar Basin[J]. Arid Zone Research, 2018, 35(5): 1146-1152. ]
[42]	苏培玺. 中国荒漠C₄木本植物和土壤无机固碳研究回顾与展望[J]. 中国沙漠, 2022, 42(1): 23-33. doi: 10.7522/j.issn.1000-694X.2021.00184
	[Su Peixi. Review and prospect of the researches on C₄ woody plants and soil inorganic carbon sequestration in deserts of China[J]. Journal of Desert Research, 2022, 42(1): 23-33. ] doi: 10.7522/j.issn.1000-694X.2021.00184

行政区	样地	经度	纬度	年均温/℃	年均降水量/mm	干旱指数	琵琶柴样本量/n	沙拐枣样本量/n
阿拉善右旗，内蒙古阿拉善盟	A	100.12°E	40.13°N	8.34	98.00	0.05	5	5
阿拉善右旗，内蒙古阿拉善盟	B	103.34°E	39.73°N	8.86	93.00	0.05	5	4
额济纳旗，内蒙古阿拉善盟	C	98.15°E	42.00°N	5.84	67.00	0.04	5	4
额济纳旗，内蒙古阿拉善盟	D	99.15°E	41.91°N	7.46	54.00	0.03	7	5
额济纳旗，内蒙古阿拉善盟	E	100.44°E	41.85°N	8.62	40.00	0.02	3	4

性状	缩写	单位
导水面积百分比	CA	%
导管密度	VD	no?mm^-2
导管成分指数	S	10⁶ mm⁴
平均导管直径	D	μm
水力直径	D_h	μm
贡献95%导水率的导管直径	D₉₅	μm
导管分组指数	V_g
导管壁厚度	t	μm
导管厚跨比	(t/b)²
木材密度	WD	g?cm^-3
理论导水率	K_th	kg?m^?1?MPa^?1?s^?1
Carlquist脆弱性指数	VI	mm?m^-2

性状	琵琶柴		沙拐枣
	D_w	D	D_w	D
CA	2.01	2	2.27	2
D	2.79	3	2.51	3
V_g	0.71	0	1.47	0
WD	1.58	1	1.18	0
(t/b)²	0.85	0	0.97	0
K_th	2.58	4	2.42	2
VI	2.25	2	1.52	1
总网络	12.76	6	12.35	4

琵琶柴和沙拐枣茎的木质部结构的差异性及空间变异特征

Differences and spatial variation in the stem xylem structural traits of Reaumuria soongarica and Calligonum mongolicum

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