全 文 :人工种植天麻的影响因子优化研究∗
张德著1ꎬ2ꎬ 美 亮1ꎬ2ꎬ 何海艳3ꎬ 王玉川3ꎬ 曹安江4ꎬ
郭 静1ꎬ2ꎬ 王咏舟1ꎬ2ꎬ 杨明挚1ꎬ 张汉波1ꎬ2∗∗
(1 云南大学生命科学院ꎬ 云南 昆明 650091ꎻ 2 云南大学生物资源保护与利用重点实验室ꎬ 云南 昆明 650091ꎻ
3 昭通天麻研究所ꎬ 云南 昭通 657000ꎻ 4 昭通林业科学研究所ꎬ 云南 昭通 657000)
摘要: 天麻是一种名贵的中草药ꎬ 可用于治疗多种疾病ꎮ 自然条件下ꎬ 天麻通过与真菌蜜环菌的共生从腐
木获取营养进行生长ꎮ 为了对人工大规模种植天麻的影响因子进行优化研究ꎬ 利用正交设计方法ꎬ 在昭通
小草坝地区开展了一个样本量较大的种植实验ꎮ 在测试的 4种因子中ꎬ 接种蜜环菌对种植天麻影响最大ꎻ
其次ꎬ 在露天灌木丛地种植的天麻产量明显高于在林地下的种植ꎮ 此外ꎬ 开展了 12 个树种对蜜环菌生长
以及天麻种植的影响研究ꎮ 结果发现十齿花 (Dipentodon sinicus) 对蜜环菌的生长最好ꎬ 同时ꎬ 也在天麻
种植实验中有最高的产量ꎻ 然而ꎬ 添加磷、 钾元素并不提高天麻的产量ꎮ
关键词: 天麻ꎻ 蜜环菌ꎻ 人工种植ꎻ 生态因子ꎻ 菌材
中图分类号: Q 948 12 文献标识码: A 文章编号: 2095-0845(2014)02-254-07
Study of Factors for Cultivating the Orchid Species
Gastrodia elataꎬ a Traditional Chinese Medicine
ZHANG De ̄Zhu1ꎬ2ꎬ MEI Liang1ꎬ2ꎬ HE Hai ̄Yan3ꎬ WANG Yu ̄Chuan3ꎬ CAO An ̄Jiang 4ꎬ
GUO Jing1ꎬ2ꎬ WANG Yong ̄Zhou1ꎬ2ꎬ YANG Ming ̄Zhi2ꎬ ZHANG Han ̄Bo1ꎬ2∗∗
(1 Laboratory of Conservation and Utilization for Bio ̄resources and Key Laboratory for Microbial Resources of the Ministry
of Educationꎬ Yunnan Universityꎬ Kunming 650091ꎬ Chinaꎻ 2 School of Life Scienceꎬ Yunnan Universityꎬ
Kunming 650091ꎬ Chinaꎻ 3 Gastrodia Tuber Research Institute of Zhaotongꎬ Zhaotong 657000ꎬ Chinaꎻ
4 Forestry Science Research Institute of Zhaotongꎬ Zhaotong 657000ꎬ China)
Abstract: The orchid speices Gastrodia elata is a valuable traditional Chinese medicine and has been widely applied
for treating a variety of diseases. The yield of wild G elata is very limited since its vegetative growth is exclusively
dependent on its symbiosis with the fungus Armillaria melleaꎬ from which G elata is able to obtain nutrients from rot ̄
ten wood in the forest. To develop a standard for cultivating G elata in large quantitiesꎬ four factors that may influ ̄
ence the yield of G elata need to be further investigatedꎬ including cultivation environmentꎬ inoculation volume of
Armillariaꎬ wood from different tree speciesꎬ and the effect of fertilizer on the maturation of G elata. To optimize
these factorsꎬ a large ̄scale orthogonal experiment was performed in the farmland of Xiaocaobaꎬ Zhaotongꎬ Yunnan
Province. Among the four factors testedꎬ inoculation of Armillaria played the most important role in the maturation of
G elata. The yield of mature tubersꎬ in terms of both tuber weight and quantityꎬ on open ground is greater than that
in forested areas. Of the 12 tree species testedꎬ Dipentodon sinicus stimulated the largest amount of growth of Armill ̄
aria and produced the greatest yield of G elata in the farmland. In comparison to the other factors testedꎬ fertilizer
showed no effect on the yield of G elata.
植 物 分 类 与 资 源 学 报 2014ꎬ 36 (2): 254~260
Plant Diversity and Resources DOI: 10.7677 / ynzwyj201413100
∗
∗∗
Funding: The National Science Foundation of China (30960077ꎬ 31360153) and Foundation of Gastrodia Tuber Research Institute of Zhaotong
Author for correspondenceꎻ E ̄mail: zhhb@ynu edu cn
Received date: 2013-04-26ꎬ Accepted date: 2013-08-20
作者简介: 张德著 (1987-) 男ꎬ 硕士研究生ꎬ 主要从事微生物生态学研究ꎮ E ̄mail: zhangdezhu1987@126 com
Key words: Gastrodia elataꎻ Armillariaꎻ Cultivationꎻ Ecological factorꎻ Wood
Gastrodiaꎬ the tuber of the orchid species Gast ̄
rodia elata Blumeꎬ is a valuable traditional Chinese
medicine and has been widely applied for treating a
variety of diseasesꎬ including headachesꎬ dizzinessꎬ
hemiplegiaꎬ rheumatismꎬ and epilepsy ( Tang and
Eisenbrandꎬ 1992). Recentlyꎬ the active compounds
from this tuber have been found to act as anticonvul ̄
sive agentsꎬ antioxidantsꎬ and antidepressants and
may also be potential treatments for Alzheimer’s dis ̄
ease (Zhou et al.ꎬ 2006ꎻ Chen et al.ꎬ 2008ꎬ 2009ꎻ
Chen and Sheenꎬ 2011). Several importantꎬ medi ̄
cally ̄relevant biological activities of Gastrodia have
been demonstratedꎬ such as anti ̄cancer ( Heo et
al.ꎬ 2007) and anti ̄asthma properties (Jang et al.ꎬ
2010). In additionꎬ Gastrodia has also been consid ̄
ered safe for long ̄term administrationꎬ and therefore
it has previously been used to improve the health and
prolong the life of humans (Chen and Sheenꎬ 2011).
In natureꎬ G elata is a perennial herb and usu ̄
ally grows on the edge of or within the forest (Zhouꎬ
1973). Because G elata is achlorophyllous and a ̄
phyllousꎬ it has to obtain nutrients by forming sym ̄
biotic relationships with two types of fungi: Mycena
osmundicola and several species from Armillaria.
While the former fungus stimulates the germination
of G elata seedꎬ the latter is thought to decompose
the wood from dead trees and thereby release into
the soil the nutrients necessary for the vegetative
growth of G elata. Only in the presence of these two
fungi can G elata complete its life cycle from seed to
protocorm to immature tuber to mature tuber (Xu et
al.ꎬ 1989). Of the Armillaria speciesꎬ A mellea is
the first reported fungus to form a symbiotic relation ̄
ship with G elata ( Kusanoꎬ 1911) and has been
considered to play the most important role in the
maturation of G elata (Anderson and Ullrichꎬ 1979ꎻ
Xu and Guoꎬ 2000ꎻ Wang et al.ꎬ 2007).
The yield of wild G elata has traditionally been
very low due to the lack of appropriate methods for
improving its cultivation. In 1968ꎬ a methodology for
cultivating G elata was invented and appliedꎬ and
since then its yield has been greatly improved. The
current cultivation process begins by first placing
several layers of fresh wood within a hole in the
ground. Naturally ̄collected tree bark and branchesꎬ
which are infected with Armillariaꎬ are then inocula ̄
ted and cultivated for nearly two months. Subse ̄
quentlyꎬ immature tubers of G elata are planted and
then cultivated after an additional year to obtain the
mature tubers (Zhang and Yangꎬ 2007).
While this cultivation process has been used for
many yearsꎬ several problems still need to be solved
in order to develop a reliable standard for cultivating
G elata in large quantities. Firstꎬ although G elata
has been commonly cultivated both in the forest and
on the open groundꎬ it is unclear if the yield of
G elata differs among these two types of environ ̄
mentsꎬ which can greatly vary in temperatureꎬ hu ̄
midityꎬ and amount of sunlight. Secondꎬ inoculating
G elata with commercial Armillaria has been widely
used to increase the yield and shorten the production
cycle of G elata (Huaꎬ 2004)ꎻ howeverꎬ the opti ̄
mal quantity of Armillaria to use for inoculations has
not been determined. Thirdꎬ Armillaria species are
able to grow on over 600 plant speciesꎬ including
treesꎬ shrubsꎬ and herbs. In practiceꎬ while farmers
usually use a mixture of wood from different tree spe ̄
cies for cultivating G elata (Zhang and Yangꎬ 2007)ꎬ
it remains to be determined how different types of
wood affects the growth of G elata. Finallyꎬ it is un ̄
clear if there are any positive effects of fertilizer on
the yield of G elata. In this studyꎬ we use an ortho ̄
gonal design to perform the first large ̄scale G elata
cultivation experiment to identify the most important
factors for improving the yieldꎬ i e. the growth and
weight of G elata.
1 Materials and methods
1 1 Description of experimental site
Cultivation of G elata was performed in Xiaocao ̄
5522期 ZHANG De ̄Zhu et al.: Study of Factors for Cultivating the Orchid Species Gastrodia elataꎬ a Traditional
baꎬ Zhaotongꎬ Yunnan Province of China ( at ap ̄
proximately 27°N and 104°Eꎬ with an average alti ̄
tude of 1 700 m). Xiaocaoba is famous for the pro ̄
duction of the highest quality of G elata specimens
in all of Chinaꎬ which is dueꎬ in partꎬ to the clima ̄
tic suitability of this area. Xiaocaoba receives an an ̄
nual rainfall of 960-1 300 mm and is characterized
by relatively high humidity (ranging from 76 to 85%
each year) (Zhang and Yangꎬ 2007). The average
temperature is above -1 ℃ in the coldest month and
is below 25 ℃ in the warmest month (Yuan et al.ꎬ
2002).
1 2 Effects of tree species on the growth of Ar ̄
millaria
In totalꎬ twelve tree speciesꎬ which are com ̄
monly used for the cultivation of G elata in Xiaocao ̄
baꎬ were selected to determine how the selection of
tree species influences the growth of Armillaria.
These tree species we chose included: Malus domes ̄
ticaꎬ Sorbus pohuashanensisꎬ Glochidion puberum
( Linn.) Hutchꎬ Dipentodon sinicus Dunnꎬ Padus
brachypoda (Batal.) Schneidꎬ Illicium verumꎬ Phoebe
zhennan S. Lee et F N. Weiꎬ Betula spꎬ Schima super ̄
ba Gardn et Champꎬ Cerasus serrula (Franch.) Yüet
Liꎬ Acer oliverianumꎬ and Lithocarpus balansae Bl.
Tree branches were freshly collected from the
local forestꎬ brought back to the laboratoryꎬ and cut
into fragments of 0 5 cm ( long) ×0 5 cm (wide) ×
0 1 cm (high). About ten grams of each wood frag ̄
ment was boiled in water for 30 minutesꎬ producing
100 mL of leaching solution. Glucose ( 2%) and
agar (1 8%) were then added to the leaching solu ̄
tion to make the medium for culturing Armillaria.
For each tree speciesꎬ we created 5 replications of
this culturing medium. After 10 days of culturing at
25 ℃ꎬ the length of the hyphae of Armillaira was
measured and used as an index of growth of the Ar ̄
millaria.
1 3 The cultivation of G elata
Typicallyꎬ it takes three years to obtain mature
tubers from the seeds of G elata. In the first yearꎬ
seeds germinate with the help of the symbiotic fungus
M osmundicola and grow into protocorms. The proto ̄
corms are then used to obtain immature tubers after
one year of cultivation with Armillaria. Finallyꎬ the
immature tubers develop into mature tubes after cul ̄
tivation with Armillaria for an additional year. Rather
than waiting for tubers to grow and mature from
seedꎬ we acquired commercially ̄developed immature
tubers that we then propagated and used to cultivate
mature tubers.
An orthogonal experiment was performed to e ̄
valuate the effects of wood typeꎬ inoculation volume
of Armillariaꎬ and fertilizers on the production of
G elata (Table 1). Each of these factors had three
levelsꎬ resulting in 9 unique combinations of factors.
Based on the results from the Armillaria growth ex ̄
perimentsꎬ only the best tree for the growth of Armil ̄
lariaꎬ Dipentodon sinicusꎬ and of the seven worst
trees for Armillaria growthꎬ only Sorbus pohuas ̄
hanensis and Betula sp.ꎬ were chosen and collected
from the local forest for the G elata cultivation ex ̄
periments. Commercially ̄produced Armillaria M1
was purchased from Zhaotong Shengnong Limitedꎬ
and we used three inoculation volumes (0ꎬ 250 and
500 g per hole) for each type of wood. In some ca ̄
sesꎬ fertilizer ( 10 g / hole) were added and com ̄
pletely mixed with soil when planting G elata.
Table 1 L9 orthogonal array applied in the cultivation of the G elata
Factor 1 2 3 4 5 6 7 8 9
Type of wood a D D D S S S B B B
Inoculation volume of Armillaria
/ ghole
0 500 250 250 0 500 500 250 0
Fertilizer K2SO4 KH2PO4 — KH2PO4 — K2SO4 — K2SO4 KH2PO4
aThe type of wood. Dꎬ Dipentodon sinicusꎻ Sꎬ Sorbus pohuashanensis and Bꎬ Betula sp
652 植 物 分 类 与 资 源 学 报 第 36卷
In September 2010ꎬ a total of 540 holesꎬ each
0 5 m (wide) ×1 m ( long) ×0 5 m (high)ꎬ were
dugꎬ half on the open ground and half in the forest.
At both of these environments (i e. ground and for ̄
est)ꎬ each orthogonal test was arranged with 9 treat ̄
ments (see above) of 30 replicates per treatment. In
each holeꎬ 5-10 freshly ̄cut wood fragments (about
10 kg in total) were laid in two layersꎬ with Armill ̄
aria M1 as needed. The holes were then filled with
soil and Armillaria was cultivated for two months. In
November 2010ꎬ the top soil of each hole was re ̄
movedꎬ and the immature tubers of G elata were
planted on top of the layers of wood. Fertilizers were
applied as neededꎬ and then we refilled the holes
with the soil. About one year laterꎬ in October 2011ꎬ
all of the mature tubers were collectedꎬ countedꎬ
and weighed.
1 4 Data analysis
All data were analyzed using the software pack ̄
age SPSS13 0 (Spssꎬ Inc.ꎬ 2004). One ̄way analy ̄
sis of variance (ANOVA) was performed to deter ̄
mine the effect of tree type on the growth of Armillar ̄
iaꎬ and compare the effects of each factor on tuber
weight in the orthogonal test previously described
(Deng et al.ꎬ 2009). The weight and number of tu ̄
bers obtained from the forest and from the open
ground was compared using a nonparametric inde ̄
pendent ̄sample test.
2 Results
2 1 Effect of wood type on the growth of Armill ̄
aria hyphae
The tree species D sinicus stimulated the grea ̄
test amount of Armillaria growth ( mean length of
3 34 cm for the rhizomorph)ꎬ which was significantly
higher than for almost all of the other tested wood types
(P < 0 05). On the other handꎬ the tree species
S pohuashanensis and Betula sp displayed the weakest
effects on the growth of Armillariaꎬ with the length of
the rhizomorphs growing less than 2 cm (Fig 1).
2 2 Cultivation of G elata
Among the four tested factorsꎬ inoculation of
Armillaria plays the most important role in determi ̄
ning the weight of the tubers ( F = 25 706ꎬ P <
0 001). The type of field environment ( i e. open
ground vs within the forest) was the second most im ̄
portant factor influencing the yield (in terms of tuber
weight) of G elata (F = 19 141ꎬ P < 0 001). In
additionꎬ the type of wood also significantly affected
the weight of tubers (F = 6 058ꎬ P = 0 003). In
contrastꎬ the addition of fertilizers had no effect on
the yield of G elata (Table 2).
Fig 1 Effects of the leaching solution of different tree woods on the growth of Armillaria.
The bars labeled with different letters are statistically significant (P < 0 05)
1. Malus domesticaꎻ 2. Sorbus pohuashanensisꎻ 3. Glochidion puberumꎻ 4. Dipentodon sinicusꎻ 5. Padus brachypodaꎻ 6. Illicium verumꎻ
7. Phoebe zhennanꎻ 8. Betula. spꎻ 9. Schima superbaꎻ 10. Cerasus serrulaꎻ 11. Acer oliverianumꎻ 12. Lithocarpus balansae
7522期 ZHANG De ̄Zhu et al.: Study of Factors for Cultivating the Orchid Species Gastrodia elataꎬ a Traditional
Table 2 Statistics of the effects of four factors on the weight of mature tubers of G elata
Source Type III Sum of Squares df Mean Square F Sig.
Inoculation volume of Armillaria 2 565 2 1 282 25 706 0 000
Field environments 0 955 1 0 955 19 141 0 000
Type of tree woods 0 604 2 0 302 6 058 0 003
Inorganic elements 0 153 2 0 077 1 534 0 217
Error 23 696 475 0 065
Total 40 398 483
Corrected Total 27 976 482
The performance of individual factors on influ ̄
encing the weight of tubers is showed in Fig 2 In ̄
oculation of Armillaria significantly affected the
weight of the tubersꎬ in an inoculation volume ̄de ̄
pendent mannerꎬ i e. greater inoculation volumes
resulted in greater tuber weights. The use of wood
from the tree species D sinicus significantly in ̄
creased the weight of the tubersꎬ nearly doubling the
Fig 2 Effect of woods from different tree species (A)ꎬ inoculation
volume of Armillaria M1 (B) and inorganic elements (C) on the
tuber weight of G elata. The bars labeled with different
letters are statistically significant (P < 0 05)
weight of the tubers when compared to using Betula
sp. (P < 0 05).
The yield of mature tubersꎬ in terms of both the
weight and the quantity producedꎬ on the open
ground is significantly greater than that from within
the forest (P < 0 05). On averageꎬ each hole from
the open ground environment yielded 3 2 tubers of
G elataꎬ weighing an average of 0 209 kgꎬ which
was nearly two times the tuber yield produced from
within the forested environment (1 8 and 0 114 kgꎬ
respectivelyꎻ Fig 3).
Fig 3 Comparison of the yield of the G elata cultivated in the
forest and on the open ground. The bars labeled with
different letters are statistically significant (P < 0 05)
3 Discussion
The vegetative growth of G elata primarily de ̄
pends upon the nutrients supplied by the decomposi ̄
852 植 物 分 类 与 资 源 学 报 第 36卷
tion of wood by the fungus Armillariaꎬ which in turn
parasitizes the plants. Thereforeꎬ the wood is a pri ̄
mary nutrient source for both G elata and Armilllaria
(Zhang and Yangꎬ 2007). Although Armillaria is a ̄
ble to infect a variety of tree species ( Sun et al.ꎬ
2007)ꎬ different trees are chemically distinctive and
could therefore differentially affect the growth of Ar ̄
millaria and consequently the production of G elata
(Rong and Caiꎬ 2010). In generalꎬ the faster and
more branched the Armillaria hyphaeꎬ the higher the
yield of G elata (Wang et al.ꎬ 2001). This study
confirmed that the type of wood does impact the
growth of Armillaria and suggested thatꎬ of the 12
tree species studiedꎬ D sinicus has the greatest
effect on increasing the hyphae growth of Armillaria
(Fig 1). Similarlyꎬ D sinicus also demonstrated a
greater ability to increase the weight of tubers in the
cultivation of G elata than the two other tree spe ̄
cies: S pohuashanensis and Betula sp. ( Fig 2 ).
Currentlyꎬ local farmers cultivate G elata using a
mixture of wood from different tree speciesꎬ primari ̄
ly selecting from among the 12 wood types used in
this study (Fig 1). This method not only causes the
low and unstable production of G elataꎬ but also re ̄
sults in the unnecessary damage of local forests re ̄
sulting from the poor utilization of appropriate wood
types. Based on our resultsꎬ we recommend that peo ̄
ple involved in the cultivation of G elata plant the
tree species D sinicusꎬ which will not only increase
the yield of G elata for the farmersꎬ but also slow
down the destruction of local forests.
The yield of tubers inoculated with the Armillar ̄
ia fungus is significantly greater than the yield with ̄
out any inoculation (Table 2). Moreoverꎬ our results
indicate that larger inoculation volumes of Armillaria
produce higher yields of G elata (Fig 2). Howeverꎬ
caution should be taken when using a large inocula ̄
tion volume of Armillaria since too much Armillaria
might negatively impact the growth of G elata
(Zhang and Yangꎬ 2007). Aside from the primary
nutrition provided by Armillariaꎬ G elata has also
been thought to be able to obtain partial secondary
nutritionꎬ e g. phosphorous uptakeꎬ from the soil
(Zhuang et al.ꎬ 1983). Howeverꎬ our results demon ̄
strated that inorganic elements have no impact on the
yield of tubers (Table 2ꎬ Fig 2). This could be be ̄
cause inorganic elements in the local soils are suffi ̄
cient for the growth requirement of G elata. Thereforeꎬ
the addition of chemical nutrients to the soils for im ̄
proving the cultivation of G elata is not recommended.
The growth of G elata depends upon optimal
lightꎬ temperatureꎬ and humidity conditions (Zhang
and Yangꎬ 2007)ꎬ which vary among forested areas
and open ground. In this experimentꎬ the total
weight of mature tubers cultivated on the open
ground was nearly two times of that produced from
the forested environmentꎬ where the temperatures
were much cooler (Fig 3)ꎬ thus supporting previous
reports that the growth of G elata benefits from high
temperatures (Liu and Wuꎬ 1998). Howeverꎬ more
immature tubers were obtained from the forest than
from the open ground (data not shown). Since these
immature tubers from the forest will take an extra
year to grow into mature tubers than those from the
open groundꎬ this implies a higher cost of time and
land ̄use for farmers if cultivating G elata from in ̄
side the forest.
Acknowledgement: We thank Yun ̄Tao Shiꎬ Yun ̄Jiao Chenꎬ
Li ̄Min Chenꎬ Yue Jiang at Yunnan Universityꎬ for partially
samplingꎬ isolating and identifying strainsꎻ Dr. Ming Zhu
from School of Medicineꎬ Yunnan Universityꎬ for critical dis ̄
cussionsꎬ commentsꎬ and language editing on the manuscript.
In particularꎬ we give our great appreciation to Dr. Steven
Callenꎬ Department of Biologyꎬ Saint Louis Universityꎬ USAꎬ
for helping us with the English language and constructive sug ̄
gestions.
Reference:
Anderson JBꎬ Ullrich RCꎬ 1979. Biological species of Armillaria mellea
in North America [J] . Mycologiaꎬ 71: 402—414
Chen PJꎬ Hsieh CLꎬ Su KP et al.ꎬ 2008. The antidepressant effect of
Gastrodia elata Bl. on the forced ̄swimming test in rats [J] . The
American Journal of Chinese Medicineꎬ 36: 95—106
Chen PJꎬ Hsieh CLꎬ Su KP et al.ꎬ 2009. Rhizomes of Gastrodia elata
Bl possess antidepressant ̄like effect via monoamine modulation
9522期 ZHANG De ̄Zhu et al.: Study of Factors for Cultivating the Orchid Species Gastrodia elataꎬ a Traditional
in subchronic animal model [J] . The American Journal of Chi ̄
nese Medicineꎬ 37: 1113—1124
Chen PJꎬ Sheen LYꎬ 2011. Gastrodiae Rhizoma (tian ma): a review
of biological activity and antidepressant mechanisms [J] . Jour ̄
nal of Traditional and Complementary Medicineꎬ 1: 31—40
Deng ZW (邓振伟)ꎬ Yu P (于萍)ꎬ Chen L (陈玲)ꎬ 2009. Appli ̄
cation of SPSS software in orthogonal design and result analysis
[J] . Computer Study (电脑学习)ꎬ 5: 15—17
Heo JCꎬ Woo SUꎬ Son M et al.ꎬ 2007. Anti ̄tumor activity of Gastro ̄
dia elata Blume is closely associated with a GTP ̄Ras ̄dependent
pathway [J] . Oncology Reportsꎬ 18: 849—853
Hua XA (华秀爱)ꎬ 2004. Rapid culture of Armillaria to promote
Gatrodia elata fast ̄growing and productive experiment [J] . Edi ̄
ble Fungi of China (中国食用菌)ꎬ 23: 27—29
Jang YWꎬ Lee JYꎬ Kim CJꎬ 2010. Anti ̄asthmatic activity of phenolic
compounds from the roots of Gastrodia elata Bl [J] . Internation ̄
al Immunopharmacologyꎬ 10: 147—154
Kusano SJꎬ 1911. Gastrodia elata and its symbiotic association with
Armillaria mellea [J] . Journal College Agricultureꎬ Imperial U ̄
niversity of Tokyoꎬ 4: 1—66
Liu NJ (刘能俊)ꎬ Wu YY (吴沿友)ꎬ 1998. Research and Applica ̄
tion of Gastrodia elata (天麻的研究与应用) [M]. Guiyang:
Guizhou Nationalities Publishing
Rong LH (容丽华)ꎬ Cai CT (蔡传涛)ꎬ 2010. Effects of different
woods on the yield of Gastrodia elata [ J] . Journal of Wuhan
Botanical Research (武汉植物学研究)ꎬ 28: 761—766
Sun LF (孙立夫)ꎬ Zhang YH (张艳华)ꎬ Yang GT (杨国亭) et
al.ꎬ 2007. Geographic distribution survey of Armillaria biological
species in Heilongjiang province [ J] . Mycosystema (菌物学
报)ꎬ 26: 59—67
Tang Wꎬ Eisenbrand Gꎬ 1992. Chinese Drugs of Plant Origin. Chem ̄
istryꎬ Pharmacologyꎬ and Use in Traditional and Modern Medi ̄
cine [M]. Berlin / Heidelberg: Springer ̄Verlag
Wang HXꎬ Yang Tꎬ Zeng Y et al.ꎬ 2007. Expression analysis of the
Gastrodianin gene ga4B in an achlorophyllous plant Gastrodia ela ̄
ta Bl. [J] . Plant Cell Reportsꎬ 26: 253—259
Wang QY (王秋颖)ꎬ Guo SX (郭顺星)ꎬ Guan FB (关凤斌)ꎬ
2001. Studies on influences of Armillaria mellea strains from dif ̄
ferent sources on yield of Gastrodia elata [J] . Chinese Tradition ̄
al and Herbal Drugs (中草药)ꎬ 32: 839—841
Xu JT (徐锦堂)ꎬ Ran YZ (冉砚珠)ꎬ Guo SX (郭顺星)ꎬ 1989.
Study on the life cycle of Gastrodia elata [ J] . Acta Academiae
Medicinae Sinicae (中国医学科学院学报)ꎬ 11: 237—241
Xu JTꎬ Guo SXꎬ 2000. Retrospect on the research of the cultivation of
Gastrodia elata Blꎬ a rare traditional Chinese medicine [J] . Chi ̄
nese Medical Journalꎬ 113: 686—692
Yuan CW (袁崇文)ꎬ Liu Z (刘智)ꎬ Yuan YQ (袁玉清) et al.ꎬ
2002. Chinese Gastrodia elata (中国天麻) [M]. Guiyang:
Guizhou Science and Technology Press
Zhang GM (张光明)ꎬ Yang LX (杨廉玺)ꎬ 2007. Research and Ex ̄
ploration of Gastrodia elata in Zhaotong (昭通天麻研究开发)
[M]. Kunming: Yunnan Science and Technology Press
Zhou BHꎬ Li XJꎬ Liu M et al.ꎬ 2006. Antidepressant ̄like activity of
the Gastrodia elata ethanol extract in mice [ J] . Fitoterapiaꎬ
77: 592—594
Zhou X (周铉)ꎬ 1973. Exploring the cultivation of the fungi ̄feeding
plant G elata [ J] . Letters in Chinese Traditional and Herbal
Drugs (ZHONGCAOYAO TONGXUN) (中草药通讯)ꎬ 2: 77
Zhuang Y (庄毅)ꎬ Wang YZ (王荣珍)ꎬ Zhang WF (张卫芳) et
al.ꎬ 1983. A study on the source of secondary nutrients for Gast ̄
rodia elata Bl. [ J] . Acta Botanica Yunnanica (云南植物研
究)ꎬ 5: 83—90
062 植 物 分 类 与 资 源 学 报 第 36卷