徐琳煜^{1, 2,},
刘守赞³,
白岩^{1, 2,,},
张汝民^{1, 2},
丁恒^{1, 2},
吴学谦^{1, 2},
郑炳松^{1, 2}
1.省部共建亚热带森林培育国家重点实验室 临安 311300
2.浙江农林大学林业与生物技术学院 临安 311300
3.浙江农林大学植物园 临安 311300
基金项目: 浙江省自然科学基金项目LY13C130010

详细信息

作者简介:徐琳煜, 主要从事药用植物栽培与生理研究。E-mail: xlyzjnl@163.com

通讯作者:白岩, 主要从事药用植物栽培与生理生化调控研究。E-mail: hzbaiyan@163.com

中图分类号:R282.2

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出版历程

收稿日期:2017-08-02
录用日期:2018-01-11
刊出日期:2018-05-01

Responses of leaf spectral characteristics of Atractylodes macrocephala Koidz. to drought stress

XU Linyu^{1, 2,},
LIU Shouzan³,
BAI Yan^{1, 2,,},
ZHANG Rumin^{1, 2},
DING Heng^{1, 2},
WU Xueqian^{1, 2},
ZHENG Bingsong^{1, 2}
1. State Key Laboratory of Subtropical Silviculture, Zhejiang Agriculture and Forestry University, Lin'an 311300, China
2. School of Forestry and Biotechnology, Zhejiang Agriculture and Forestry University, Lin'an 311300, China
3. Botanical Garden, Zhejiang Agricultural and Forestry University, Lin'an 311300, China
Funds: the Natural Science Foundation of Zhejiang Province of ChinaLY13C130010

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Corresponding author:BAI Yan,E-mail: hzbaiyan@163.com

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摘要
摘要:白术根茎膨大初期对轻度的土壤水分缺失具有一定的耐受性，但是过度干旱会抑制其根茎膨大及成分积累，因此，无损伤诊断该时期白术是否受到干旱胁迫至关重要。本文以2年生白术为试验材料，在根茎膨大初期控制土壤水分，形成不同程度的干旱胁迫，利用UniSpec-SC光谱分析仪测定其光谱反射率，并结合光合色素含量，探讨白术叶片光谱特征对干旱胁迫的响应规律，为利用光谱参数监测白术生长状况提供技术依据。结果显示，随着干旱胁迫程度增加，可见光区域（400～750 nm）白术叶片光谱反射率升高，说明其光能吸收利用能力下降；但在750～1 000 nm的近红外波段，光谱反射率则逐渐平稳，在1 000 nm处，干旱胁迫下的全部叶片反射率均低于对照。微分光谱在680～750 nm间差异明显，并与叶绿素含量在700～750 nm间呈显著相关。同时，大多数光谱参数与色素含量呈显著相关（P < 0.05），尤其类胡萝卜指数（mCRI）、色素归一化指数（PSNDb）、红边位置（λ_red）、红边幅值（D_λred）、红边面积（S_red）与之呈极显著相关（P < 0.01）。从上述结果可知，微分光谱680～750 nm可作为检测白术是否受到干旱影响的主要监测波段，红边参数、类胡萝卜指数及色素归一化指数可以作为重要指标，快速、准确、无伤害诊断白术受干旱胁迫程度。本研究结果可为应用反射光谱进行白术干旱胁迫程度诊断提供理论依据和技术支持。
关键词:白术/
干旱胁迫/
光谱特征/
光谱反射率/
光合色素/
光谱参数
Abstract:Atractylodes macrocephala Koidz. is a perennial herb belonging to Compositae family, which is fond of cool climate regions. Rhizome dried for over 2 years are used for a series of medicinal functions. It is a top medicinal herb in Zhejiang Province, China. At the early stage of rhizome enlargement, A. macrocephala has a certain tolerance to mild soil drought, but excessive drought can inhibit rhizome enlargement and accumulation of constituent chemicals. Increasing degree of drought stress could slow down growth, inhibit rhizome enlargement and limit yield. In order to provide reference for drought stress control and cultivation of drought-resistant varieties, biennial A. macrocephala were planted under different drought stress. Spectral reflectance of A. macrocephala leaves determined by UniSpec-SC spectrum analyzer and combined with photosynthetic pigment contents were used to explain the response of spectral characteristics under drought stress. The results showed that spectral reflectance increased in the visible region (400-750 nm) with increasing drought stress. This indicated that the absorption and utilization ability of light energy decreased under increased drought stress. However, spectral reflectance gradually stabilized in the near-infrared band 750-1 000 nm. The reflectance of all the leaves under drought stress was lower than that of the control at 1 000 nm. The difference in spectrum of 680-750 nm was significant, which was correlated with chlorophyll content in 700-750 nm. This band could be used to monitor whether A. macrocephala was affected by drought. The contents of photosynthetic pigments increased initially and then decreased with the increased drought stress. It indicated that a suitable degree of drought was good for growth. Most of the spectral parameters were significantly correlated with pigments contents (P < 0.05). Spectral parameters of mCRI, PSNDb, red-edge position (λ_red), red-edge amplitude (D_λred) and red-edge area (S_red) were significantly correlated with leaf pigments contents (P < 0.01), which could be used to diagnose drought indicators. In summary, differential spectrum of 680-750 nm could be used to detect drought impact on A. macrocephala. Red-edge parameters, carotenoid reflectance indexes and pigment specific normalized difference may be used as indicators to diagnose drought stress degree of A. macrocephala. This conclusion not only provided a reference for the study of high-spectrum plant research, but also provided theoretical basis and technical support for the application of spectrum diagnosis of A. macrocephala in drought stress analysis.
Key words:Atractylodes macrocephala Koidz./
Drought stress/
Spectral characteristics/
Spectral reflectance/
Photosynthetic pigment/
Spectral parameter

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图1不同干旱胁迫对白术叶片反射光谱的影响
CK:对照组, 正常浇水, 即每天浇水量0.5 L; Tr1、Tr2、Tr3和Tr4为干旱处理组, 分别停止浇水2 d、4 d、6 d和8 d。
Figure1.Effect of drought stress on reflectance spectra of Atractylodes macrocephala leaves
CK: control group, watered 0.5 L per day; Tr1, Tr2, Tr3 and Tr4 were drought-treated groups, stopped watering for 2 days, 4 days, 6 days and 8 days, respectively.


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图2不同干旱胁迫处理白术叶片反射光谱一阶导数
CK:对照组, 正常浇水, 即每天浇水量0.5 L; Tr1、Tr2、Tr3和Tr4为干旱处理组, 分别停止浇水2 d、4 d、6 d和8 d。
Figure2.First derivative of reflectance spectra of Atractylodes macrocephala leaves under different drought stresses
CK: control group, watered 0.5 L per day; Tr1, Tr2, Tr3 and Tr4 were drought-treated groups, stopped watering for 2 days, 4 days, 6 days and 8 days, respectively.


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图3白术叶片光合色素含量与光谱反射率的相关性
Ca:叶绿素a; Cb:叶绿素b; Cars:胡萝卜素; a/b:叶绿素a/b; T:总叶绿素含量; 0.05:显著性水平为0.05; 0.01:显著性水平为0.01。
Figure3.Correlation between reflectance spectra and photosynthetic pigments contents of Atractylodes macrocephala leaves
Ca: chlorophyll a; Cb: chlorophyll b; Cars: carotene; a/b: chlorophyll a/b; T: total chlorophyll content; 0.05: significant level 0.05; 0.01: significant level 0.01.


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图4白术叶片光合色素含量与微分光谱的相关性
Ca:叶绿素a; Cb:叶绿素b; Cars:胡萝卜素; a/b:叶绿素a/b; T:总叶绿素含量; 0.05:显著性水平为0.05; 0.01:显著性水平为0.01。
Figure4.Correlation between derivative spectra and photosynthetic pigments contents of Atractylodes macrocephala leaves
Ca: chlorophyll a; Cb: chlorophyll b; Cars: carotene; a/b: chlorophyll a/b; T: total chlorophyll content; 0.05: significant level 0.05; 0.01: significant level 0.01.


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表1不同水分胁迫处理的土壤含水量
Table1.Soil water contents of different water stress treatments

%
	CK	Tr1	Tr2	Tr3	Tr4
土壤含水量Soil water content	52.52±1.07a	44.37±1.07b	31.40±1.15c	21.97±0.55d	12.70±0.60d
CK:对照组，正常浇水，每天浇水量0.5 L; Tr1、Tr2、Tr3和Tr4为干旱处理组，分别停止浇水2d、4d、6d和8d。不同字母表示在0.05水平上差异显著。CK: control group, watered 0.5 L per day; Tr1, Tr2, Tr3 and Tr4 were drought-treated groups, stopped watering for 2 days, 4 days, 6 days and 8 days, respectively. Different lowercase letters mean significant differences at 0.05 level.

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表2干旱胁迫对白术叶片光合色素含量的影响
Table2.Effect of drought stress on photosynthetic pigments contents of Atractylodes macrocephala leaves

光合色素Photosynthetic pigment	CK	Tr1	Tr2	Tr3	Tr4
叶绿素a (Chla) Chlorophyll a (mg·g-1)	2.115±0.132a	2.352±0.211a	2.707±0.362a	2.053±0.484a	0.893±0.018b
叶绿素b (Chlb) Chlorophyll b (mg·g-1)	0.923±0.034ab	1.017±0.063a	1.016±0.123b	0.762±0.153a	0.420±0.027c
叶绿素总量(Chla+b) Total chlorophyll (mg·g-1)	3.038±0.166a	3.369±0.273a	3.723±0.486a	2.815±0.636a	1.312±0.044b
类胡萝卜素Carotenoids (mg·g-1)	0.330±0.006bc	0.482±0.061a	0.470±0.099bc	0.292±0.092ab	0.173±0.021c
叶绿素a/b (Chl a/b) Chlorophyll a/b	2.290±0.060b	2.311±0.063b	2.662±0.036a	2.681±0.146a	2.132±0.102b
CK:对照组，正常浇水，每天浇水量0.5 L; Tr1、Tr2、Tr3和Tr4为干旱处理组，分别停止浇水2 d、4 d、6 d和8 d。同行不同字母表示在0.05水平上差异显著。CK: control group, watered 0.5 L per day; Tr1, Tr2, Tr3 and Tr4 were drought-treated groups, stopped watering for 2 days, 4 days, 6 days and 8 days, respectively. Different lowercase letters in the same line mean significant differences at 0.05 level.

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表3不同干旱胁迫处理下白术叶片红边、黄边和蓝边参数
Table3.Red edge, yellow edge and blue edge parameters of Atractylodes macrocephala leaves under different drought stressess

参数Parameter	CK	Tr1	Tr2	Tr3	Tr4
红边位置(λred) Red edge position	704.667±0.557a	714.333±1.528a	705.000±1.000a	708.000±5.196a	707.333±9.292a
红边幅值（Dλred) Red edge amplitude	1.346±0.076a	1.154±0.059ab	1.263±0.142ab	1.080±0.062b	1.053±0.085b
红边面积(Sred) Red edge area	51.281±1.477ab	53.687±2.436ab	58.431±3.204a	54.132±4.092ab	47.161±4.665b
黄边位置(λyellow) Yellow edge position	571.667±1.528a	570.000±0.000a	570.667±1.155a	571.333±2.309a	571.667±2.082a
黄边幅值(Dλyellow)Yellow edge amplitude	0.182±0.057a	0.243±0.008a	0.242±0.043a	0.242±0.025a	0.230±0.018a
黄边面积(Syellow) Yellow edge area	-8.757±0.649a	-10.002±0.404a	-9.972±1.430a	-10.575±1.486a	-9.860±1.086a
蓝边位置(λblue) Blue edge position	520.333±0.577a	520.333±3.780a	521.333±0.577a	518.000±5.000a	519.000±2.646a
蓝边幅值(Dλblue) Blue edge amplitude	0.375±0.0278a	0.394±0.016a	0.417±0.062a	0.438±0.070a	0.408±0.050a
蓝边面积(Sblue) Blue edge area	9.427±0.796a	10.366±0.654a	11.057±1.586a	11.768±1.855a	10.827±1.302a
CK:对照组，正常浇水，即每天浇水量0.5 L; Tr1、Tr2、Tr3和Tr4为干旱处理组，分别停止浇水2 d、4 d、6 d和8 d。同行不同字母表示在0.05水平上差异显著。CK: control group，watered 0.5 L per day; Tr1，Tr2, Tr3 and Tr4 were drought-treated groups，stopped watering for 2 days，4 days，6 days and 8 days，respectively. Different lowercase letters in the same line mean significant differences at 0.05 level.

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表4不同干旱胁迫处理下白术叶片反射光谱参数的影响s
Table4.Effect of drought stress on reflectance spectra parameter values of Atractylodes macrocephala leaves

参数Parameter	CK	Tr1	Tr2	Tr3	Tr4
反射率倒数(RR) Reciprocal reflectance	4.622±0.430a	4.841±0.297a	4.478±0.414ab	4.188±0.485ab	3.504±0.054b
反射光谱指数680 (SR680) Spectral reflectance index 680	7.649±1.378a	7.259±0.691a	7.428±0.402a	6.624±0.032a	4.326±0.353b
反射光谱指数705 (SR705) Spectral reflectance index 705	2.100±0.082b	2.399±0.179a	2.180±0.106ab	2.180±0.070ab	1.716±0.027c
反射光谱比值指数a (RARSa) Reflectance spectrum ratio index a	0.368±0.042ab	0.386±0.025ab	0.358±0.004b	0.383±0.022ab	0.444±0.043a
反射光谱比值指数b (RARSb) Reflectance spectrum ratio index b	0.451±0.052ab	0.452±0.010ab	0.434±0.010b	0.527±0.006a	0.521±0.038a
归一化指数(NDVI) Normalized difference vegetation index	0.765±0.040a	0.757±0.021a	0.762±0.011a	0.738±0.001a	0.623±0.025b
红度归一化指数(rNDVI) Red edge NDVI	0.355±0.017b	0.411±0.032a	0.371±0.021ab	0.371±0.014ab	0.263±0.007c
改良的红边比值指数(mND705) Modified red edge ratio index 705	0.454±0.010a	0.523±0.041a	0.466±0.024a	0.472±0.028a	0.380±0.024b
绿度归一化指数(gNDN) Green normalized difference	0.493±0.040a	0.489±0.030a	0.484±0.019a	0.432±0.021a	0.353±0.003b
色素归一化差值指数a (PSNDa) Pigment specific normalized difference a	0.774±0.039a	0.774±0.020a	0.778±0.010a	0.754±0.002a	0.641±0.026b
色素归一化差值指数b (PSNDb) Pigment specific normalized difference b	0.730±0.044a	0.740±0.025a	0.737±0.010a	0.676±0.010a	0.591±0.021b
结构不敏感色素指数(SIPI) Structure-insensitive pigment	0.747±0.032a	0.746±0.020a	0.753±0.009a	0.738±0.007a	0.615±0.024b
类胡萝卜素指数1 (CRI1) Carotenoid reflectance index 1	3.567±0.842ab	3.376±0.253ab	3.817±0.278a	2.557±0.212bc	2.011±0.221c
类胡萝卜素指数2 (CRI2) Carotenoid reflectance index 2	3.919±1.028a	3.223±0.193ab	3.971±0.407a	2.343±0.252b	2.114±0.231b
改良类胡萝卜指数(mCRI) Modified carotenoid reflectance index	2.201±0.498ab	2.131±0.228ab	2.430±0.116a	1.665±0.041bc	1.167±0.139c
光化学反射指数(PRI) Photochemical reflectance index	0.016±0.004b	0.033±0.003a	0.012±0.007b	0.018±0.003b	0.016±0.001b
红绿比指数(RGI) Red/green index	0.676±0.025ab	0.642±0.013b	0.663±0.016b	0.665±0.014b	0.723±0.026a
CK:对照组, 正常浇水, 即每天浇水量0.5 L; Tr1、Tr2、Tr3和Tr4为干旱处理组, 分别停止浇水2 d、4 d、6 d和8 d。同行不同字母表示在0.05水平上差异显著。CK: control group, watered 0.5 L per day; Tr1, Tr2, Tr3 and Tr4 were drought-treated groups, stopped watering for 2 days, 4 days, 6 days and 8 days, respectively. Different lowercase letters in the same line mean significant differences at 0.05 level.

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表5不同干旱胁迫处理下白术叶片反射光谱参数与光合色素含量的相关性
Table5.Correlation between reflectance spectra parameters and photosynthetic pigments contents of Atractylodes macrocephala leaves under different drought stresses

参数 Parameter	叶绿素a Chlorophyll a	叶绿素b Chlorophyll b	叶绿素a+b Chlorophyll a+b	类胡萝卜素 Carotinoid
反射率倒数(RR) Reciprocal reflectance	0.868	0.962**	0.900*	0.873
反射光谱指数680 (SR680) Spectral reflectance index 680	0.931*	0.960**	0.946*	0.807
反射光谱指数705 (SR705) Spectral reflectance index 705	0.866	0.893*	0.880*	0.872
反射光谱比值指数a (RARSa) Reflectance spectrum ratio index a	-0.948*	-0.900*	-0.944*	-0.757
反射光谱比值指数b (RARSb) Reflectance spectrum ratio index b	-0.742	-0.837	-0.773	-0.827
归—化指数(NDVI) Normalized difference vegetation index	0.945*	0.955*	0.954*	0.809
红度归一化指数(rNDVI) Red edge NDVI	0.891*	0.908*	0.902*	0.867
改良的红边比值指数(mND705) Modified red edge ratio index 705	0.820	0.860	0.837	0.850
绿度归一化指数(gNDN) Green normalized difference	0.915*	0.980**	0.939*	0.861
色素归一化差值指数a (PSNDa) Pigment specific normalized difference a	0.956*	0.961**	0.965**	0.830
色素归一化差值指数b (PSNDb) Pigment specific normalized difference b	0.943*	0.995**	0.964**	0.905*
结构不敏感色素指数(SIPI) Structure-insensitive pigment	0.952*	0.939*	0.955*	0.801
类胡萝卜素指数1 (CRI1) Carotenoid reflectance index 1	0.892*	0.940*	0.911*	0.863
类胡萝卜素指数2 (CRI2) Carotenoid reflectance index 2	0.764	0.828	0.787	0.722
改良类胡萝卜指数(mCRI) Modified carotenoid reflectance index	0.936*	0.964**	0.950*	0.888*
光化学反射指数(PRI) Photochemical reflectance index	0.131	0.287	0.174	0.394
红绿比指数(RGI) Red/green index	-0.894*	-0.905*	-0.904*	-0.881*
红边位置(λred) Red edge position	0.045	0.151	0.074	0.328
红边幅值(Dλred) Red edge amplitude	0.604	0.685	0.630	0.492
红边面积(Sred) Red edge area	0.924*	0.796	0.896*	0.816
*表示相关性显著(P < 0.05); **表示相关性极显著(P < 0.01)。* and ** mean significant correlation at 0.05 and 0.01 levels, respectively.

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