Morphological traits and physiological characteristics in drought tolerance in 20 shrub species on the Qinghai-Xizang Plateau

doi:10.3724/SP.J.1258.2014.00052

Abstract

Abstract:

Aims Providing indispensably theoretical evidence for establishing indices of evaluation on drought tolerance in shrubs, and screening for tree species that are drought tolerant for afforestation in arid regions or for matching their characteristics with suitable habitat conditions are the key to vegetation restoration in the Qinghai-Xizang Plateau. However, these issues are not adequately addressed in recent research due to lack of systematic methods. Therefore, our objective was to make a comprehensive evaluation on drought tolerance in 20 shrub species collected from different areas in Qinghai-Xizang Plateau, and to study their underlying mechanisms in drought tolerance.
Methods We made measurements on variables depicting root characteristics, including the root length (TRL), surface area (TRSA), volume (TRV), and tips number (TRTN) of all roots, the root length (FRL), surface area (FRSA), volume (FRV), and tips number (FRTN) of fine roots (d≤2 mm), and derived plant characteristic indices including thickness of cuticle (CT), thickness of palisade tissue (TPT), thickness of spongy tissue (TST), TPT/TST, thickness of leaf (LT), palisable tissue cell density, and tissue structural tense ratio (CTR = TPT/LT × 100%) and spongy tissue loosened ratio (SR = TST/LT × 100%) of leaf anatomical structure, root to shoot ratio (RSR), leaf transpiration rate (T_r), instantaneous water use efficiency (WUE_i), and carbon isotopic composition (δ¹³C) of the 20 shrub species through field experiments. Correlation analysis and principal component analysis were performed on the 19 variables and indices.
Important findings Different shrubs had different mechanisms of drought tolerance. In this study, the character- istics of drought tolerance were mainly categorized into 6 types, involving modifications of (1) root systems, (2) leaf anatomical structure, (3) leaf pattern, and (4) biomass allocation, or via (5) low water-consumption and (6) high WUE_i. Different genera or different tree species within the same genus clearly differed in drought tolerance. The species of the genus Hippophae were relatively poorly tolerant to drought, whereas several shrubs including Potentilla fruticosa, Berberis julianae, Caragana arborescens, Spiraea salicifolia and Hippophae rhamnoides ssp. mongolica occurred to be more drought tolerant than other shrub species investigated in this study. On the other hand, there were highly significant correlations among the characteristics of root systems and among characteristics of leaf anatomical structure. The results of principal component analysis on 19 variables and indices showed that TRL, TRSA, TRV, TRTN, FRL, FRSA, FRV, FRTN, CT, TPT, TST and WUE_i could be effective indicators of drought tolerance of shrubs in the Qinghai-Xizang Plateau. In addition, the drought tolerance of shrubs had a close connection with their origin of collections; the shrubs collected from Xining prefecture in Qinghai Province were more drought tolerant than those from Tianshui Prefecture in Gansu and Lasa Prefecture in Xizang.

Key words: characteristics of root system, drought resistance, leaf anatomical structure, Qinghai-Xizang Plateau, shrub

QIU Quan,PAN Xin,LI Ji-Yue,WANG Jun-Hui,MA Jian-Wei,DU Kun. Morphological traits and physiological characteristics in drought tolerance in 20 shrub species on the Qinghai-Xizang Plateau[J]. Chin J Plant Ecol, 2014, 38(6): 562-575.

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URL: https://www.plant-ecology.com/EN/10.3724/SP.J.1258.2014.00052

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Figures/Tables 9

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树种 Tree species	缩写 Abbreviation	采集地点 Collection site	经纬度 Longitude & latitude	海拔 Altitude (m)
柠条锦鸡儿 Caragana korshinskii	CK	青海西宁南山 Nanshan, Xining, Qinghai	101.27° E 36.43° N	2 545
短叶锦鸡儿 Caragana brevifolia	CB	青海大通县宝库乡 Baoku Township, Datong County, Qinghai	101.35° E 37.12° N	3 200
尼泊尔锦鸡儿 Caragana sukiensis	CSU	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
锦鸡儿 Caragana sinica	CSI	青海省农林科学院 Qinghai Academy of Agricultural and Forestry	101.45° E 36.43° N	2 285
树锦鸡儿 Caragana arborescens	CA	青海省农林科学院 Qinghai Academy of Agricultural and Forestry	101.45° E 36.43° N	2 285
蒙古沙棘 Hippophae rhamnoides ssp. mongolica	HRR	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
江孜沙棘 Hippophae rhamnoides subsp. gyantsensis	HRG	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
柳叶沙棘 Hippophae salicifolia	HS	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
西藏沙棘 Hippophae thibetana	HT	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
豪猪刺 Berberis julianae	BJ	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
西北小檗 Berberis vernae	BV	青海门源县仙米林场 Xianmi Forest Center, Menyuan County, Qinghai	102.01° E 37.15° N	2 730
金露梅 Potentilla fruticosa	PF	青海门源县仙米林场 Xianmi Forest Center, Menyuan County, Qinghai	102.00° E 37.18° N	2 817
野蔷薇 Rosa multiflora	RM	青海大通县宝库乡 Baoku Township, Datong County, Qinghai	101.35° E 37.12° N	3 036
胡颓子 Elaeagnus pungens	EP	甘肃小陇山林业科学研究所 Xiaolongshan Forestry Science and Technology Research Institute, Gansu	105.80° E 34.48° N	1 450
光果莸 Caryopteris tangutica	CT	青海西宁北山 Beishan, Xining, Qinghai	101.45° E 36.44° N	2 482
驼绒藜 Ceratoides latens	CL	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
鲜卑花 Sibiraea laevigata	SL	青海大通县宝库乡 Baoku Township, Datong County, Qinghai	101.36° E 37.13° N	3 202
绣线菊 Spiraea salicifoli	SS	甘肃小陇山林业科学研究所 Xiaolongshan Forestry Science and Technology Research Institute, Gansu	105.80° E 34.48° N	1 450
西北沼委陵菜 Comarum salesovianum	CS	青海门源县仙米林场 Xianmi Forest Center, Menyuan County, Qinghai	102.58° E 37.24° N	3 081
华北珍珠梅 Sorbaria kirilowii	SK	甘肃小陇山林业科学研究所 Xiaolongshan Forestry Science and Technology Research Institute, Gansu	105.80° E 34.48° N	1 450

树种 Tree species	缩写 Abbreviation	采集地点 Collection site	经纬度 Longitude & latitude	海拔 Altitude (m)
柠条锦鸡儿 Caragana korshinskii	CK	青海西宁南山 Nanshan, Xining, Qinghai	101.27° E 36.43° N	2 545
短叶锦鸡儿 Caragana brevifolia	CB	青海大通县宝库乡 Baoku Township, Datong County, Qinghai	101.35° E 37.12° N	3 200
尼泊尔锦鸡儿 Caragana sukiensis	CSU	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
锦鸡儿 Caragana sinica	CSI	青海省农林科学院 Qinghai Academy of Agricultural and Forestry	101.45° E 36.43° N	2 285
树锦鸡儿 Caragana arborescens	CA	青海省农林科学院 Qinghai Academy of Agricultural and Forestry	101.45° E 36.43° N	2 285
蒙古沙棘 Hippophae rhamnoides ssp. mongolica	HRR	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
江孜沙棘 Hippophae rhamnoides subsp. gyantsensis	HRG	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
柳叶沙棘 Hippophae salicifolia	HS	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
西藏沙棘 Hippophae thibetana	HT	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
豪猪刺 Berberis julianae	BJ	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
西北小檗 Berberis vernae	BV	青海门源县仙米林场 Xianmi Forest Center, Menyuan County, Qinghai	102.01° E 37.15° N	2 730
金露梅 Potentilla fruticosa	PF	青海门源县仙米林场 Xianmi Forest Center, Menyuan County, Qinghai	102.00° E 37.18° N	2 817
野蔷薇 Rosa multiflora	RM	青海大通县宝库乡 Baoku Township, Datong County, Qinghai	101.35° E 37.12° N	3 036
胡颓子 Elaeagnus pungens	EP	甘肃小陇山林业科学研究所 Xiaolongshan Forestry Science and Technology Research Institute, Gansu	105.80° E 34.48° N	1 450
光果莸 Caryopteris tangutica	CT	青海西宁北山 Beishan, Xining, Qinghai	101.45° E 36.44° N	2 482
驼绒藜 Ceratoides latens	CL	西藏堆龙德庆县柳梧乡 Liuwu Township, Doilungdêqên County, Xizang	91.03° E 29.57° N	3 630
鲜卑花 Sibiraea laevigata	SL	青海大通县宝库乡 Baoku Township, Datong County, Qinghai	101.36° E 37.13° N	3 202
绣线菊 Spiraea salicifoli	SS	甘肃小陇山林业科学研究所 Xiaolongshan Forestry Science and Technology Research Institute, Gansu	105.80° E 34.48° N	1 450
西北沼委陵菜 Comarum salesovianum	CS	青海门源县仙米林场 Xianmi Forest Center, Menyuan County, Qinghai	102.58° E 37.24° N	3 081
华北珍珠梅 Sorbaria kirilowii	SK	甘肃小陇山林业科学研究所 Xiaolongshan Forestry Science and Technology Research Institute, Gansu	105.80° E 34.48° N	1 450

树种缩写 Tree species abbreviation	TRL (cm)	TRSA (cm²)	TRV (cm³)	TRTN	FRL (cm)	FRSA (cm²)	FRV (cm³)	FRTN	RSR	T_r (mmol·m^-2·s^-1)
CK	2 091.47 ± 57.03	960.63 ± 114.23	74.10 ± 12.36	4 022 ± 99	1 704.75 ± 98.83	357.74 ± 13.50	9.27 ± 0.24	3 002 ± 92	0.50 ± 0.09	3.98 ± 0.09
CB	795.19 ± 97.95	497.44 ± 7.94	31.73 ± 0.09	2 473 ± 37	576.30 ± 45.16	138.45 ± 14.02	3.96 ± 0.15	1 757 ± 50	0.40 ± 0.07	3.57 ± 0.03
CSU	506.70 ± 56.20	109.36 ± 12.73	3.98 ± 0.66	1 203 ± 385	457.23 ± 24.14	81.18 ± 3.57	1.79 ± 0.10	1 149 ± 32	0.80 ± 0.20	3.80 ± 0.22
CSI	1 084.17 ± 102.42	412.54 ± 13.69	36.24 ± 3.51	2 364 ± 172	892.11 ± 102.94	203.62 ± 22.48	4.82 ± 0.94	1 833 ± 122	0.23 ± 0.04	3.53 ± 0.10
CA	4 067.11 ± 92.60	1 204.68 ± 58.43	71.05 ± 5.80	8 365 ± 798	3 651.55 ± 154.64	759.97 ± 21.12	18.32 ± 0.25	6 757 ± 571	0.33 ± 0.04	4.98 ± 0.21
HRR	1 047.89 ± 84.08	425.90 ± 36.68	37.33 ± 2.00	1 944 ± 141	868.55 ± 11.42	199.40 ± 7.74	5.46 ± 0.67	1 488 ± 139	0.14 ± 0.03	4.69 ± 0.06
HRG	282.74 ± 32.93	80.11 ± 2.39	4.01 ± 0.17	718 ± 25	255.68 ± 33.43	50.49 ± 3.38	1.24 ± 0.09	567 ± 155	0.12 ± 0.05	4.84 ± 0.06
HS	518.83 ± 23.15	128.91 ± 3.84	5.97 ± 0.62	1 224 ± 163	488.65 ± 27.51	90.32 ± 3.32	2.15 ± 0.05	864 ± 67	0.11 ± 0.01	4.47 ± 0.24
HT	720.44 ± 77.80	330.29 ± 31.69	25.28 ± 1.62	1 750 ± 74	545.78 ± 63.51	133.12 ± 17.56	3.71 ± 0.46	1 242 ± 68	0.23 ± 0.04	4.30 ± 0.55
BJ	5 656.55 ± 309.05	1 213.39 ± 131.00	120.48 ± 17.40	9 631 ± 337	5 371.61 ± 402.26	972.90 ± 73.67	20.92 ± 1.65	8 522 ± 895	0.20 ± 0.01	3.57 ± 0.33
BV	777.70 ± 48.24	160.10 ± 20.52	5.79 ± 0.93	1 991 ± 189	713.83 ± 73.61	121.56 ± 18.43	2.48 ± 0.48	1 410 ± 246	0.47 ± 0.05	5.11 ± 0.35
PF	6 241.53 ± 379.90	1 797.66 ± 71.20	148.54 ± 16.52	17 139 ± 1925	4 972.53 ± 152.21	936.41 ± 123.89	23.86 ± 2.82	12 516 ± 795	0.56 ± 0.14	3.24 ± 0.06
RM	2 618.31 ± 261.83	1 003.94 ± 100.39	105.70 ± 7.57	7 780 ± 778	2 232.63 ± 223.26	415.07 ± 41.51	9.31 ± 0.93	5 851 ± 585	0.66 ± 0.14	4.84 ± 0.37
EP	2 435.75 ± 106.87	1 052.54 ± 49.77	101.16 ± 8.21	4 150 ± 44	2 217.08 ± 87.74	451.15 ± 19.41	11.44 ± 0.26	3 318 ± 54	0.72 ± 0.10	3.67 ± 0.12
CT	1 654.42 ± 66.57	728.20 ± 86.52	46.95 ± 4.34	4 331 ± 319	1 202.48 ± 235.46	275.77 ± 19.36	7.92 ± 0.59	3 027 ± 248	0.25 ± 0.03	3.91 ± 0.14
CL	444.56 ± 28.47	194.11 ± 6.13	10.22 ± 1.78	1 237 ± 99	401.59 ± 23.81	77.66 ± 9.27	1.78 ± 0.43	788 ± 64	0.36 ± 0.01	3.64 ± 0.21
SL	1 175.41 ± 131.21	321.41 ± 40.25	16.65 ± 1.65	2 908 ± 210	1 051.13 ± 153.00	201.16 ± 43.11	4.64 ± 1.26	2 255 ± 213	0.62 ± 0.16	5.84 ± 0.56
SS	4 090.83 ± 793.40	1 231.59 ± 73.07	109.77 ± 2.97	11 461 ± 821	3 114.82 ± 335.05	537.87 ± 42.75	12.32 ± 2.01	7 742 ± 618	0.51 ± 0.11	4.29 ± 0.64
CS	492.29 ± 32.28	142.85 ± 6.63	8.93 ± 0.24	1 149 ± 87	367.73 ± 67.02	63.18 ± 5.06	1.54 ± 0.10	754 ± 29	0.29 ± 0.10	6.61 ± 0.83
SK	1 852.45 ± 110.20	553.12 ± 11.11	43.15 ± 3.20	3 796 ± 196	1 733.97 ± 191.18	300.90 ± 13.50	7.24 ± 0.31	2 977 ± 130	0.50 ± 0.06	3.75 ± 0.43
平均值 Average	1 927.72	621.20	50.35	5 536	1 575.04	302.62	7.28	3 324	0.40	4.33
标准偏差 Standard deviation	1 772.54	494.73	45.22	6 282	1 464.50	265.61	6.19	3 153	0.21	0.86
变异系数 Coefficient of variation (%)	91.95	79.64	89.81	113.48	92.98	87.77	85.03	94.85	51.62	19.81

树种缩写 Tree species abbreviation	TRL (cm)	TRSA (cm²)	TRV (cm³)	TRTN	FRL (cm)	FRSA (cm²)	FRV (cm³)	FRTN	RSR	T_r (mmol·m^-2·s^-1)
CK	2 091.47 ± 57.03	960.63 ± 114.23	74.10 ± 12.36	4 022 ± 99	1 704.75 ± 98.83	357.74 ± 13.50	9.27 ± 0.24	3 002 ± 92	0.50 ± 0.09	3.98 ± 0.09
CB	795.19 ± 97.95	497.44 ± 7.94	31.73 ± 0.09	2 473 ± 37	576.30 ± 45.16	138.45 ± 14.02	3.96 ± 0.15	1 757 ± 50	0.40 ± 0.07	3.57 ± 0.03
CSU	506.70 ± 56.20	109.36 ± 12.73	3.98 ± 0.66	1 203 ± 385	457.23 ± 24.14	81.18 ± 3.57	1.79 ± 0.10	1 149 ± 32	0.80 ± 0.20	3.80 ± 0.22
CSI	1 084.17 ± 102.42	412.54 ± 13.69	36.24 ± 3.51	2 364 ± 172	892.11 ± 102.94	203.62 ± 22.48	4.82 ± 0.94	1 833 ± 122	0.23 ± 0.04	3.53 ± 0.10
CA	4 067.11 ± 92.60	1 204.68 ± 58.43	71.05 ± 5.80	8 365 ± 798	3 651.55 ± 154.64	759.97 ± 21.12	18.32 ± 0.25	6 757 ± 571	0.33 ± 0.04	4.98 ± 0.21
HRR	1 047.89 ± 84.08	425.90 ± 36.68	37.33 ± 2.00	1 944 ± 141	868.55 ± 11.42	199.40 ± 7.74	5.46 ± 0.67	1 488 ± 139	0.14 ± 0.03	4.69 ± 0.06
HRG	282.74 ± 32.93	80.11 ± 2.39	4.01 ± 0.17	718 ± 25	255.68 ± 33.43	50.49 ± 3.38	1.24 ± 0.09	567 ± 155	0.12 ± 0.05	4.84 ± 0.06
HS	518.83 ± 23.15	128.91 ± 3.84	5.97 ± 0.62	1 224 ± 163	488.65 ± 27.51	90.32 ± 3.32	2.15 ± 0.05	864 ± 67	0.11 ± 0.01	4.47 ± 0.24
HT	720.44 ± 77.80	330.29 ± 31.69	25.28 ± 1.62	1 750 ± 74	545.78 ± 63.51	133.12 ± 17.56	3.71 ± 0.46	1 242 ± 68	0.23 ± 0.04	4.30 ± 0.55
BJ	5 656.55 ± 309.05	1 213.39 ± 131.00	120.48 ± 17.40	9 631 ± 337	5 371.61 ± 402.26	972.90 ± 73.67	20.92 ± 1.65	8 522 ± 895	0.20 ± 0.01	3.57 ± 0.33
BV	777.70 ± 48.24	160.10 ± 20.52	5.79 ± 0.93	1 991 ± 189	713.83 ± 73.61	121.56 ± 18.43	2.48 ± 0.48	1 410 ± 246	0.47 ± 0.05	5.11 ± 0.35
PF	6 241.53 ± 379.90	1 797.66 ± 71.20	148.54 ± 16.52	17 139 ± 1925	4 972.53 ± 152.21	936.41 ± 123.89	23.86 ± 2.82	12 516 ± 795	0.56 ± 0.14	3.24 ± 0.06
RM	2 618.31 ± 261.83	1 003.94 ± 100.39	105.70 ± 7.57	7 780 ± 778	2 232.63 ± 223.26	415.07 ± 41.51	9.31 ± 0.93	5 851 ± 585	0.66 ± 0.14	4.84 ± 0.37
EP	2 435.75 ± 106.87	1 052.54 ± 49.77	101.16 ± 8.21	4 150 ± 44	2 217.08 ± 87.74	451.15 ± 19.41	11.44 ± 0.26	3 318 ± 54	0.72 ± 0.10	3.67 ± 0.12
CT	1 654.42 ± 66.57	728.20 ± 86.52	46.95 ± 4.34	4 331 ± 319	1 202.48 ± 235.46	275.77 ± 19.36	7.92 ± 0.59	3 027 ± 248	0.25 ± 0.03	3.91 ± 0.14
CL	444.56 ± 28.47	194.11 ± 6.13	10.22 ± 1.78	1 237 ± 99	401.59 ± 23.81	77.66 ± 9.27	1.78 ± 0.43	788 ± 64	0.36 ± 0.01	3.64 ± 0.21
SL	1 175.41 ± 131.21	321.41 ± 40.25	16.65 ± 1.65	2 908 ± 210	1 051.13 ± 153.00	201.16 ± 43.11	4.64 ± 1.26	2 255 ± 213	0.62 ± 0.16	5.84 ± 0.56
SS	4 090.83 ± 793.40	1 231.59 ± 73.07	109.77 ± 2.97	11 461 ± 821	3 114.82 ± 335.05	537.87 ± 42.75	12.32 ± 2.01	7 742 ± 618	0.51 ± 0.11	4.29 ± 0.64
CS	492.29 ± 32.28	142.85 ± 6.63	8.93 ± 0.24	1 149 ± 87	367.73 ± 67.02	63.18 ± 5.06	1.54 ± 0.10	754 ± 29	0.29 ± 0.10	6.61 ± 0.83
SK	1 852.45 ± 110.20	553.12 ± 11.11	43.15 ± 3.20	3 796 ± 196	1 733.97 ± 191.18	300.90 ± 13.50	7.24 ± 0.31	2 977 ± 130	0.50 ± 0.06	3.75 ± 0.43
平均值 Average	1 927.72	621.20	50.35	5 536	1 575.04	302.62	7.28	3 324	0.40	4.33
标准偏差 Standard deviation	1 772.54	494.73	45.22	6 282	1 464.50	265.61	6.19	3 153	0.21	0.86
变异系数 Coefficient of variation (%)	91.95	79.64	89.81	113.48	92.98	87.77	85.03	94.85	51.62	19.81

树种 Tree species	LT (μm)	CT (μm)	TPT (μm)	TST (μm)	TPT/TST	CTR (%)	SR (%)	δ¹³C (‰)	WUE_i (μmol·mmol^-1)
CK	200.39 ± 12.39	1.03 ± 0.27	75.94 ± 5.60	106.18 ± 13.00	0.73 ± 0.1	38.06 ± 2.89	52.97 ± 3.84	-30.22 ± 0.47	2.38 ± 0.20
CB	107.63 ± 13.01	2.70 ± 1.16	40.20 ± 10.91	44.28 ± 2.51	0.91 ± 0.25	36.75 ± 6.56	41.68 ± 5.21	-28.40 ± 0.44	3.57 ± 0.05
CSU	138.33 ± 6.92	4.83 ± 1.56	50.81 ± 5.32	56.37 ± 4.67	0.91 ± 0.13	36.59 ± 3.54	40.59 ± 2.84	-28.06 ± 0.44	3.34 ± 0.01
CSI	150.36 ± 14.74	3.15 ± 0.83	77.35 ± 11.47	37.21 ± 6.74	2.13 ± 0.44	52.56 ± 4.48	25.29 ± 3.47	-28.86 ± 0.45	3.04 ± 0.12
CA	130.36 ± 13.27	1.15 ± 0.45	72.57 ± 10.49	35.57 ± 9.93	2.22 ± 0.75	55.03 ± 6.11	26.8 ± 6.48	-28.61 ± 0.45	2.53 ± 0.02
HRR	299.65 ± 4.64	0.93 ± 0.50	165.71 ± 3.27	104.02 ± 2.43	1.59 ± 0.04	55.64 ± 0.61	34.93 ± 0.58	-28.15 ± 0.44	3.23 ± 0.07
HRG	116.3 ± 13.17	2.81 ± 0.47	39.62 ± 6.07	51.81 ± 9.72	0.78 ± 0.17	33.87 ± 4.02	44.08 ± 4.58	-30.22 ± 0.47	1.85 ± 0.05
HS	120.72 ± 23.68	1.09 ± 0.26	41.71 ± 6.9	52.83 ± 24.15	0.98 ± 0.52	36.13 ± 8.49	42.4 ± 11.73	-28.71 ± 0.45	3.19 ± 0.09
HT	132.82 ± 15.47	2.56 ± 0.38	56.29 ± 7.82	41.96 ± 10.75	1.46 ± 0.51	42.10 ± 3.73	30.86 ± 6.06	-27.68 ± 0.43	3.70 ± 0.08
BJ	246.30 ± 8.62	2.11 ± 0.29	114.6 ± 6.93	105.31 ± 8.02	1.10 ± 0.12	46.59 ± 2.56	42.78 ± 2.49	-27.14 ± 0.42	4.27 ± 0.03
BV	244.01 ± 6.73	0.78 ± 0.3	86.71 ± 4.53	125.58 ± 3.89	0.69 ± 0.05	36.11 ± 1.69	52.31 ± 1.52	-28.69 ± 0.45	2.86 ± 0.02
PF	154.23 ± 9.43	2.65 ± 0.94	69.62 ± 4.87	45.00 ± 5.25	1.57 ± 0.2	45.63 ± 2.4	29.47 ± 2.99	-26.09 ± 0.41	4.43 ± 0.09
RM	126.32 ± 5.87	1.66 ± 0.22	57.66 ± 2.59	47.73 ± 3.64	1.21 ± 0.1	46.36 ± 1.96	38.31 ± 1.6	-27.46 ± 0.43	3.83 ± 0.01
EP	205.98 ± 15.56	0.9 ± 0.24	104.76 ± 3.21	78.25 ± 14.90	1.74 ± 2.4	51.17 ± 5.18	37.54 ± 6.4	-28.92 ± 0.45	2.82 ± 0.05
CT	248.30 ± 14.84	1.14 ± 0.35	61.41 ± 13.41	152.00 ± 7.83	0.4 ± 0.09	24.68 ± 5.09	61.83 ± 4.15	-28.60 ± 0.45	2.72 ± 0.19
CL	186.24 ± 32.07	1.72 ± 0.22	78.57.03 ± 15.12	79.51 ± 8.12	0.99 ± 0.08	42.19 ± 3.48	42.69 ± 3.94	-28.36 ± 0.44	3.23 ± 0.17
SL	135.45 ± 18.49	0.96 ± 0.30	53.00 ± 12.08	60.36 ± 5.38	0.87 ± 0.14	38.38 ± 3.88	44.56 ± 3.23	-29.85 ± 0.47	2.35 ± 0.09
SS	213.32 ± 12.19	1.24 ± 0.22	72.81 ± 4.77	113.58 ± 9.48	0.64 ± 0.06	33.76 ± 1.73	52.57 ± 2.33	-29.28 ± 0.46	2.43 ± 0.01
CS	163.33 ± 6.13	0.94 ± 0.39	87.21 ± 3.19	48.36 ± 4.11	1.82 ± 0.17	54.32 ± 1.98	30.08 ± 2.01	-28.56 ± 0.45	2.64 ± 0.12
SK	183.95 ± 13.66	3.11 ± 0.77	95.88 ± 7.16	55.98 ± 9.84	1.77 ± 0.35	52.75 ± 3.08	30.63 ± 4.33	-29.11 ± 0.45	2.51 ± 0.32
平均值 Average	175.20	1.87	75.12	72.09	1.23	42.93	40.12	175.20	1.87
标准偏差 Standard deviation	66.28	1.08	29.86	33.95	0.53	8.72	9.79	66.28	1.08
变异系数 Coefficient of variation (%)	37.83	57.69	39.75	47.09	42.86	20.30	24.40	37.83	57.69