沙冬青根瘤菌的抗逆性

doi:10.17521/cjpe.2006.0020

摘要/Abstract

摘要：

沙冬青(Ammopiptanthus mongolicus)是西北荒漠地区珍稀常绿阔叶灌木,综合沙冬青分布区内气候、土壤和水分等因子,选取宁夏中卫县沙坡头、内蒙古磴口县、内蒙古阿拉善左旗、内蒙古乌拉特后旗作为样区,于2004年4月中下旬沙冬青的花期调查和收集根瘤,并结合地理环境对分离得到的根瘤菌进行抗逆性初步分析。调查发现根瘤的生成及其在宿主植物上的着生部位、大小、形状、颜色等因生境条件的不同而有所差异,水分是其主要的影响因子;分离得到根瘤菌17株,对其耐盐性、耐酸碱性、生长温度范围和抗生素抗性进行了研究。结果表明,64.7%的菌株可以在含3%NaCl的YMA培养基上生长;94.1%的菌株可以在pH 5~11的范围内生长;全部菌株在60 ℃处理10 min后仍能生长;不同菌株对不同抗生素表现出不同的抗性,ZW₄和WH₄¹对各抗生素表现出较强的抗性。分离自磴口县的根瘤菌普遍表现出较强的抗酸碱和抗高温的能力,这是对其环境的适应。本研究证明沙冬青根瘤菌具有较强的抗逆性,但菌株间存在差异,这可能是对沙冬青分布区多种景观生态类型的适应。同时测定了几种伴生植物中间锦鸡儿(Caragana intermedia)、猫头刺(Oxytropis aciphylla)和柠条(Caragana corshinskii)根瘤菌的抗逆性,与沙冬青根瘤菌表现出一定的相似性,具较强的抗逆性,说明生态环境对豆科植物-根瘤菌共生体有重要的影响。沙冬青的高抗性可能与沙冬青根瘤菌的高抗性有关。沙冬青与其根瘤菌的共生关系有利于其植物群落的发育和多样性的维系。

关键词: 沙冬青, 根瘤菌, 抗逆性, 生态环境, 高抗性

Abstract:

Ammopiptanthus mongolicus is an endangered evergreen broad-leaf species in the northwest desert zone of China. Shapotou, Alashanzuoqi, Dengkou and Wulatehouqi were defined as research regions based on soil, water and other ecological factors. Nodules of A. mongolicus were collected during the last twenty-days of April, which was the flowering period of the host, and their characteristics were studied. The resistance of rhizobia isolated from A. mongolicus also was studied in relation to environmental factors of the distribution areas. Seventeen rhizobia strains were isolated from A. mongolicus. It was found that nodules had various attachment modes, sizes, shapes and colors, which were related to differences of their environment, of which water may be the most important determinant. Several biochemical characteristics were determined, including resistance to salt, acid-alkali, temperature variation and intrinsic antibiotics. The results indicated that 64.7% strains could tolerate NaCl stress at 3% concentration, 94.1% strains could grow at a range of pH from 5-11, and all strains could grow after being exposed to 60 ℃ for 10 min. Differences in resistance to different intrinsic antibiotics existed among strain ZW₄, and WH₄¹ had high a resistance to different intrinsic antibiotics. Rhizobia strains from Dengkou had higher resistance to acid-alkali conditions and temperature. These results indicated that rhizobia strains isolated from A. mongolicus were highly resistant to stresses. However, there were differences among strains in their resistance to stresses, which might be related to adaptations of rhizobia to the diverse landscapes in different regions. Resistances of rhizobia strains isolated from accompanying plants, for example, Caragana corshinskii, Caragana intermedia and Oxytropis aciphylla, also were studied, and the results indicated that they also were highly resistant to stresses similar to the rhizobia strains isolated from A. mongolicus. These results indicated that the symbiosis between rhizobia and legumes was influenced by their environment. The high resistance of A. mongolicus may be related to the high resistance of its rhizobia, and that the symbiotic relation was beneficial to the development and maintenance of its community in diverse landscapes.

Key words: Ammopiptanthus mongolicus, Rhizobia, Resistance, Eco-environment, High resistance

何恒斌, 贾昆峰, 贾桂霞, 丁琼. 沙冬青根瘤菌的抗逆性. 植物生态学报, 2006, 30(1): 140-146. DOI: 10.17521/cjpe.2006.0020

HE Heng-Bin, JIA Kun-Feng, JIA Gui-Xia, DING Qiong. A PRELIMINARY STUDY ON THE STRESS RESISTANCE OF RHIZOBIA ISOLATED FROM AMMOPIPTANTHUS MONGOLICUS. Chinese Journal of Plant Ecology, 2006, 30(1): 140-146. DOI: 10.17521/cjpe.2006.0020

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2006.0020

https://www.plant-ecology.com/CN/Y2006/V30/I1/140

图/表 5

表1 试验菌株采集地点及参比菌株来源

Table 1 Sampling places of the experimental strains and the source of the reference strain

菌株号Strains number	宿主名Host	采集地点Sampling places	地形及土壤Terrain and soil
$DK 1 1$	沙冬青 Ammopiptanthus mongolicus	磴口Dengkou	平坦风蚀沙地,风沙土加粘土Wind erosion moving sand, windy sand soil and clay
$DK 1 2$	沙冬青A. mongolicus	磴口Dengkou	平坦风蚀沙地,风沙土加粘土Wind erosion moving sand, windy sand soil and clay
DK₄	沙冬青A. mongolicus	磴口Dengkou	盐渍化滩地,粗砾质风沙土Saline-alkaline land, windy stone-sand soil
DK₅	沙冬青A. mongolicus	磴口Dengkou	洪水冲积沟滩地,粉尘风沙土Land caused by seasonal floods, windy sand soil
DK₆	沙冬青A. mongolicus	磴口Dengkou	洪水冲积沟滩地,粉尘风沙土Land caused by seasonal floods, windy sand soil
DK₈	沙冬青A. mongolicus	磴口Dengkou	盐渍化滩地,粗砾质风沙土Saline-alkaline land, windy stone-sand soil
ZW₄	沙冬青A. mongolicus	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
ZW₇	沙冬青A. mongolicus	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
ZW₈	沙冬青A. mongolicus	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
$WH 1 1$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 1 2$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 1 3$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 4 1$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 4 2$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
WH₆	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
AZ₁	沙冬青A. mongolicus	阿拉善左旗Alashanzuoqi	低山残塬,红土质灰棕漠土Loess hill, grey brown desert soil
AZ₂	沙冬青A. mongolicus	阿拉善左旗Alashanzuoqi	残存河床,风沙土Riverbed, windy sand soil
CAN(参比菌株Reference strain)	沙冬青A. mongolicus	磴口Dengkou	购于中国农业大学Bought from China Agriculture Universituy
WHn	柠条Caragana korshinskii	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
WHj	锦鸡儿C. intermedia	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
ZWn	柠条C. korshinskii	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
WHm	猫头刺Oxytropis aciphylla	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil

表1 试验菌株采集地点及参比菌株来源

Table 1 Sampling places of the experimental strains and the source of the reference strain

菌株号Strains number	宿主名Host	采集地点Sampling places	地形及土壤Terrain and soil
$DK 1 1$	沙冬青 Ammopiptanthus mongolicus	磴口Dengkou	平坦风蚀沙地,风沙土加粘土Wind erosion moving sand, windy sand soil and clay
$DK 1 2$	沙冬青A. mongolicus	磴口Dengkou	平坦风蚀沙地,风沙土加粘土Wind erosion moving sand, windy sand soil and clay
DK₄	沙冬青A. mongolicus	磴口Dengkou	盐渍化滩地,粗砾质风沙土Saline-alkaline land, windy stone-sand soil
DK₅	沙冬青A. mongolicus	磴口Dengkou	洪水冲积沟滩地,粉尘风沙土Land caused by seasonal floods, windy sand soil
DK₆	沙冬青A. mongolicus	磴口Dengkou	洪水冲积沟滩地,粉尘风沙土Land caused by seasonal floods, windy sand soil
DK₈	沙冬青A. mongolicus	磴口Dengkou	盐渍化滩地,粗砾质风沙土Saline-alkaline land, windy stone-sand soil
ZW₄	沙冬青A. mongolicus	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
ZW₇	沙冬青A. mongolicus	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
ZW₈	沙冬青A. mongolicus	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
$WH 1 1$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 1 2$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 1 3$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 4 1$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
$WH 4 2$	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
WH₆	沙冬青A. mongolicus	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
AZ₁	沙冬青A. mongolicus	阿拉善左旗Alashanzuoqi	低山残塬,红土质灰棕漠土Loess hill, grey brown desert soil
AZ₂	沙冬青A. mongolicus	阿拉善左旗Alashanzuoqi	残存河床,风沙土Riverbed, windy sand soil
CAN(参比菌株Reference strain)	沙冬青A. mongolicus	磴口Dengkou	购于中国农业大学Bought from China Agriculture Universituy
WHn	柠条Caragana korshinskii	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
WHj	锦鸡儿C. intermedia	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil
ZWn	柠条C. korshinskii	中卫沙坡头Shapotou	低山坡地,红土质灰棕漠土Hillside, grey brown desert soil
WHm	猫头刺Oxytropis aciphylla	乌拉特后旗Wulatehouqi	平坦石质滩地及沙丘,风沙土和红土质灰棕漠土Flood land and dune, windy sand soil and grey brown desert soil

表2 不同浓度的NaCl对根瘤菌生长的影响

Table 2 Effects of NaCl on growth of rhizobium

菌株 Strains	NaCl (%)
菌株 Strains	1	2	3	4	5	6	7
$DK 1 1$	+	+	+	-	-	-	-
$DK 1 2$	+	+	-	-	-	-	-
DK₄	+	+	+	+	+	+	-
DK₅	+	-	-	-	-	-	-
DK₆	+	-	-	-	-	-	-
DK₈	+	+	+	+	+	+	+
ZW₄	+	+	-	-	-	-	-
ZW₇	+	+	+	+	+	-	-
ZW₈	+	+	+	+	+	+	+
$WH 1 1$	+	-	-	-	-	-	-
$WH 1 2$	+	-	-	-	-	-	-
$WH 1 3$	-	-	-	-	-	-	-
$WH 4 1$	+	-	-	-	-	-	-
$WH 4 2$	+	+	+	+	-	-	-
WH₆	+	+	+	-	-	-	-
AZ₁	-	-	-	-	-	-	-
AZ₂	+	+	+	+	+	+	+
CAN	+	+	+	-	-	-	-
WHn	+	+	+	+	+	+	-
WHj	+	+	+	+	+	+	-
ZWn	-	-	-	-	-	-	-
WHm	+	+	+	+	+	+	+

表2 不同浓度的NaCl对根瘤菌生长的影响

Table 2 Effects of NaCl on growth of rhizobium

菌株 Strains	NaCl (%)
菌株 Strains	1	2	3	4	5	6	7
$DK 1 1$	+	+	+	-	-	-	-
$DK 1 2$	+	+	-	-	-	-	-
DK₄	+	+	+	+	+	+	-
DK₅	+	-	-	-	-	-	-
DK₆	+	-	-	-	-	-	-
DK₈	+	+	+	+	+	+	+
ZW₄	+	+	-	-	-	-	-
ZW₇	+	+	+	+	+	-	-
ZW₈	+	+	+	+	+	+	+
$WH 1 1$	+	-	-	-	-	-	-
$WH 1 2$	+	-	-	-	-	-	-
$WH 1 3$	-	-	-	-	-	-	-
$WH 4 1$	+	-	-	-	-	-	-
$WH 4 2$	+	+	+	+	-	-	-
WH₆	+	+	+	-	-	-	-
AZ₁	-	-	-	-	-	-	-
AZ₂	+	+	+	+	+	+	+
CAN	+	+	+	-	-	-	-
WHn	+	+	+	+	+	+	-
WHj	+	+	+	+	+	+	-
ZWn	-	-	-	-	-	-	-
WHm	+	+	+	+	+	+	+

表3 不同的pH值对根瘤菌生长的影响

Table 3 Effects of pH on growth of rhizobium

菌株 Strains	pH
菌株 Strains	3	4	5	6	8	9	10	11	12	13
$DK 1 1$	-	+	+	+	+	+	+	+	-	-
$DK 1 2$	-	+	+	+	+	+	+	+	-	-
DK₄	-	-	-	+	+	+	+	+	+	-
DK₅	-	-	+	+	+	+	+	+	-	-
DK₆	-	+	+	+	+	+	+	+	+	-
DK₈	-	+	+	+	+	+	+	+	+	-
ZW₄	-	-	+	+	+	+	+	+	+	-
ZW₇	-	-	+	+	+	+	+	+	-	-
ZW₈	-	-	+	+	+	+	+	+	+	-
$WH 1 1$	-	-	+	+	+	+	+	+	-	-
$WH 1 2$	-	+	+	+	+	+	+	+	-	-
$WH 1 3$	-	+	+	+	+	+	+	+	-	-
$WH 4 1$	-	+	+	+	+	+	+	+	-	-
$WH 4 2$	-	-	+	+	+	+	+	+	+	-
WH₆	-	-	+	+	+	+	+	+	-	-
AZ₁	-	-	+	+	+	+	+	+	-	-
AZ₂	-	+	+	+	+	+	+	+	+	-
CAN	-	-	-	-	+	+	+	+	-	-
WHn	-	-	-	-	+	+	+	+	+	-
WHj	-	+	+	+	+	+	+	+	+	-
ZWn	-	-	+	+	+	+	+	+	-	-
WHm	-	+	+	+	+	+	+	+	+	-

表3 不同的pH值对根瘤菌生长的影响

Table 3 Effects of pH on growth of rhizobium

菌株 Strains	pH
菌株 Strains	3	4	5	6	8	9	10	11	12	13
$DK 1 1$	-	+	+	+	+	+	+	+	-	-
$DK 1 2$	-	+	+	+	+	+	+	+	-	-
DK₄	-	-	-	+	+	+	+	+	+	-
DK₅	-	-	+	+	+	+	+	+	-	-
DK₆	-	+	+	+	+	+	+	+	+	-
DK₈	-	+	+	+	+	+	+	+	+	-
ZW₄	-	-	+	+	+	+	+	+	+	-
ZW₇	-	-	+	+	+	+	+	+	-	-
ZW₈	-	-	+	+	+	+	+	+	+	-
$WH 1 1$	-	-	+	+	+	+	+	+	-	-
$WH 1 2$	-	+	+	+	+	+	+	+	-	-
$WH 1 3$	-	+	+	+	+	+	+	+	-	-
$WH 4 1$	-	+	+	+	+	+	+	+	-	-
$WH 4 2$	-	-	+	+	+	+	+	+	+	-
WH₆	-	-	+	+	+	+	+	+	-	-
AZ₁	-	-	+	+	+	+	+	+	-	-
AZ₂	-	+	+	+	+	+	+	+	+	-
CAN	-	-	-	-	+	+	+	+	-	-
WHn	-	-	-	-	+	+	+	+	+	-
WHj	-	+	+	+	+	+	+	+	+	-
ZWn	-	-	+	+	+	+	+	+	-	-
WHm	-	+	+	+	+	+	+	+	+	-

表4 温度对根瘤菌生长的影响

Table 4 Effects of temperature on growth of rhizobium

菌株 Strains	温度 Temperature (℃)
菌株 Strains	4	12	20	37	45	50(10 min)	60(10 min)
$DK 1 1$	-	+	+	+	-	+	+
$DK 1 2$	-	+	+	+	-	+	+
DK₄	+	+	+	+	+	+	+
DK₅	+	+	+	-	-	+	+
DK₆	-	+	+	+	+	+	+
DK₈	-	+	+	+	+	+	+
ZW₄	+	+	+	+	-	+	+
ZW₇	-	+	+	+	-	+	+
ZW₈	+	+	+	+	+	+	+
$WH 1 1$	-	+	+	+	-	+	+
$WH 1 2$	-	+	+	+	-	+	+
$WH 1 3$	-	-	+	+	-	+	+
$WH 4 1$	-	+	+	+	-	+	+
$WH 4 2$	-	+	+	+	-	+	+
WH₆	-	+	+	+	-	+	+
AZ₁	+	+	+	+	-	+	+
AZ₂	+	+	+	+	-	+	+
CAN	+	+	+	+	-	+	+
WHn	-	+	+	+	-	+	+
WHj	-	+	+	+	-	+	+
ZWn	-	-	+	-	-	+	+
WHm	-	-	+	+	+	+	+

表4 温度对根瘤菌生长的影响

Table 4 Effects of temperature on growth of rhizobium

菌株 Strains	温度 Temperature (℃)
菌株 Strains	4	12	20	37	45	50(10 min)	60(10 min)
$DK 1 1$	-	+	+	+	-	+	+
$DK 1 2$	-	+	+	+	-	+	+
DK₄	+	+	+	+	+	+	+
DK₅	+	+	+	-	-	+	+
DK₆	-	+	+	+	+	+	+
DK₈	-	+	+	+	+	+	+
ZW₄	+	+	+	+	-	+	+
ZW₇	-	+	+	+	-	+	+
ZW₈	+	+	+	+	+	+	+
$WH 1 1$	-	+	+	+	-	+	+
$WH 1 2$	-	+	+	+	-	+	+
$WH 1 3$	-	-	+	+	-	+	+
$WH 4 1$	-	+	+	+	-	+	+
$WH 4 2$	-	+	+	+	-	+	+
WH₆	-	+	+	+	-	+	+
AZ₁	+	+	+	+	-	+	+
AZ₂	+	+	+	+	-	+	+
CAN	+	+	+	+	-	+	+
WHn	-	+	+	+	-	+	+
WHj	-	+	+	+	-	+	+
ZWn	-	-	+	-	-	+	+
WHm	-	-	+	+	+	+	+

表5 沙冬青根瘤菌对抗生素的抗性

Table 5 The resistance of Ammopiptanthus mongolicus thtzobia to antibioties

菌株 Strains	Km(μg·ml^-1)				Amp(μg·ml^-1)				Rif(μg·ml^-1)				Cm(μg·ml^-1)
菌株 Strains	5	50	100	300	5	50	100	300	5	50	100	300	5	50	100	300
$DK 1 1$	+	+	-	-	+	+	+	+	+	-	-	-	+	+	+	+
$DK 1 2$	+	+	+	-	+	+	+	+	+	-	-	-	+	+	+	+
DK₄	-	-	-	-	+	-	-	-	-	-	-	-	+	-	-	-
DK₅	+	+	-	-	+	+	+	-	-	-	-	-	+	+	-
DK₆	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-
DK₈	-	-	-	-	+	-	-	-	-	-	-	-	-	-	-	-
ZW₄	+	+	+	-	+	+	+	+	+	+	+	-	+	+	+	+
ZW₇	+	-	-	-	-	-	-	-	-	-	-	-	-	-	-	-
ZW₈	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-
$WH 1 1$	-	-	-	-	+	-	-	-	+	-	-	-	+	+	+	+
$WH 1 2$	+	-	-	-	+	-	-	-	-	-	-	-	+	+	+	+
$WH 1 3$	+	+	-	-	+	+	+	+	-	-	-	-	+	+	+	+
$WH 4 1$	+	+	+	-	+	+	+	+	+	+	-	-	+	+	+	+
$WH 4 2$	-	-	-	-	+	+	+	+	-	-	-	-	+	-	-	-
WH₆	+	+	-	-	+	+	+	+	+	-	-	-	+	+	+	-
AZ₁	+	+	-	-	+	-	-	-	-	-	-	-	+	-	-	-
AZ₂	+	-	-	-	+	+	+	+	+	-	-	-	+	+	+	+
CAN	+	-	-	-	+	+	+	+	-	-	-	-	+	+	+	+
WHn	-	-	-	-	+	-	-	-	-	-	-	-	+	-	-	-
WHj	+	-	-	-	+	+	-	-	-	-	-	-	+	+	+	+
ZWn	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-
WHm	-	-	-	-	+	-	-	-	-	-	-	-	-	-	-	-

表5 沙冬青根瘤菌对抗生素的抗性

Table 5 The resistance of Ammopiptanthus mongolicus thtzobia to antibioties

菌株 Strains	Km(μg·ml^-1)				Amp(μg·ml^-1)				Rif(μg·ml^-1)				Cm(μg·ml^-1)
菌株 Strains	5	50	100	300	5	50	100	300	5	50	100	300	5	50	100	300
$DK 1 1$	+	+	-	-	+	+	+	+	+	-	-	-	+	+	+	+
$DK 1 2$	+	+	+	-	+	+	+	+	+	-	-	-	+	+	+	+
DK₄	-	-	-	-	+	-	-	-	-	-	-	-	+	-	-	-
DK₅	+	+	-	-	+	+	+	-	-	-	-	-	+	+	-
DK₆	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-
DK₈	-	-	-	-	+	-	-	-	-	-	-	-	-	-	-	-
ZW₄	+	+	+	-	+	+	+	+	+	+	+	-	+	+	+	+
ZW₇	+	-	-	-	-	-	-	-	-	-	-	-	-	-	-	-
ZW₈	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-
$WH 1 1$	-	-	-	-	+	-	-	-	+	-	-	-	+	+	+	+
$WH 1 2$	+	-	-	-	+	-	-	-	-	-	-	-	+	+	+	+
$WH 1 3$	+	+	-	-	+	+	+	+	-	-	-	-	+	+	+	+
$WH 4 1$	+	+	+	-	+	+	+	+	+	+	-	-	+	+	+	+
$WH 4 2$	-	-	-	-	+	+	+	+	-	-	-	-	+	-	-	-
WH₆	+	+	-	-	+	+	+	+	+	-	-	-	+	+	+	-
AZ₁	+	+	-	-	+	-	-	-	-	-	-	-	+	-	-	-
AZ₂	+	-	-	-	+	+	+	+	+	-	-	-	+	+	+	+
CAN	+	-	-	-	+	+	+	+	-	-	-	-	+	+	+	+
WHn	-	-	-	-	+	-	-	-	-	-	-	-	+	-	-	-
WHj	+	-	-	-	+	+	-	-	-	-	-	-	+	+	+	+
ZWn	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-
WHm	-	-	-	-	+	-	-	-	-	-	-	-	-	-	-	-

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