全 文 ：Biological Characters of Uncinula adunca
Parasitizing in Introduced Plants of the Genus Salix
Tomoshevich M.A.
(Central Siberian Botanical Garden , SB RAS , Novosibirsk 630090 , Russia)
Abstract:Biological characters of Uncinula adunca which is a pathogene of powdery mildew of willows
are introduced.Differences in symptoms , formation of anamorphic and teleomorphic stages of the Siberian
fungus population are revealed.Some morphological characters of the pathogene carposomes are indicated.
Key words:Salix L.;powdery mildew;Uncinula adunca
CLC number:Q949.327.6 Document code:A Article ID:1672-3538(2007)04-0233-03
寄生于柳属植物的 Uncinula adunca
的生物学特性
Tomoshevich M.A.
(俄罗斯科学院 西伯利亚中心植物园 , 新西伯利亚 630090 , 俄罗斯)
摘 要:介绍了柳树白粉病病原菌 Uncinula adunca 的生物学特性 , 揭示了西伯利亚地区该菌在症状 、无性型
阶段及有性型阶段形成等方面的不同 ,同时还指出了该病原菌子实体的一些生物学特性。
关键词:柳属;白粉病;Uncinula adunca
中图分类号:Q949.327.6 文献标识码:A 文章编号:1672-3538(2007)04-0233-04
The genus Salix comprises 300—400 species dis-
tributed mainly in the Holarctic.Willows represent the
largest genus of the family Salicaceae Mirbel in the ar-
boriflora of Siberia.Willows are found everywhere as
wide thickets , they are characterized by high plastici-
ty , which allows them to adapt to various geographical
and ecological conditions , and by ability to grow well
on overmoistened soils.Due to these characteristics
willows are promising for regeneration , urban plantings
and landscaping.In dendrology and forestry , when in-
troducing woody plants , revealing pathogens and study
of their biology provide a basis for development of mea-
sures intended to plant protection from different disea
ses.Data on the mycobiota of introduced plants must
be always taken into account when evaluating potential
needs for practical use.
Data on pathogenic mycobiota of wild and intro-
duced species of the genus Salix are limited.Accord-
ing to the literature diseases causing damage of leaves
(powdery mildew , rust and spottiness)and shoot dry-
ing out are the most injurious.Most researchers agree
that powdery mildew is a dominant disease of leaves.
Powdery mildews parasitize in a great diversity of
plants.They were recorded world-wide in the plants
belonging to 88 families.Four pathogenic agents of this
disease:Podosphaera schlechtendalii Lev., Uncinula
adunca (Wallr.:Fr.)Lev.[ U.salicis(DC).Wint.
Rabenh.] , U.salici-gracilistylae Homma and Phyl-
lactinia salicis (Jacz.)Bunk were described for Salix
species.The most abundant among them was the sec-
ond pathogenic agent[ 1-4].The genus Uncinula com-
bines a range of high-organized species parasitizing
mainly in woody plants.Development of mycelium and
conidial sporulation as cobweb-like , filmy or tomentose
Biography:Tomoshevich M.А.(1975-), female , studying on the biodiversity of pathogenic micromycetes of Siberian woody plants.
Received date:2007-04-05
Corresponding author:Tomoshevich M.А., E-mai l:tomoshevich@csbg.nsc.ru
第 5卷第 4 期 菌 物 研 究 Vol.5 No.4
2007 年 12月 Journal of Fungal Research Dec.2007
farina on the upper leaf surface is typical of this genus.
Mycelium is kept safe.Cleistothecia are globe-shaped ,
often with numerous appendages.
Mycelium and conidial sporulation as filmy farina
or tomentose spots on both leaf surfaces are characteris-
tic of U.adunca.Cleistothecia are scattered or in
groups , often radially arranged , and with a great quan-
tity of appendages (60—120 and more).Some au-
thors[ 4] characterize this species as a complex one
which was recorded in 3 genera of the family Sali-
caceae:Salix , Populus and Chosenia.Sizes of cleis-
tothecia and asci and number of ascospores vary with
the genus of a host plant.
It has been established that the pathogen para-
sitizes in S.acutifolia Wild., S.alba L., S.auri-
ta L., S.brachypoda Kom., S.bebbiana Sarg., S.
caprea L., S.cinerea L., S.myrsinifolia Salisb.,
S.myrtilloides L., S.rosmarinifolia L., S.rorida
Laksch., S. schwerinii E.Wolf., S.viminalis L.
and other species[ 3-4].
1 Materials and Methods
Different species of willows introduced in the ar-
boretum of Central Siberian Botanical Garden , SB RAS
were the subjects of study.The collection is represen-
ted by 60 Salix species united in 148 groups differing
in ecological and geographical characters.Study of
powdery mildew was begun in 1997.
2 Results and discussion
It was established that , on the average , up to
25% of plant groups were affected by powdery mildew
(Uncinula adunca).Annually this disease is recorded
in S.pyrolifolia Ledeb., S.lanata L., S.saxatilis
Turcz. ex Ledeb., S. taraikensis Kimura , S.
caprea , S.myrtilloides , S.rorida , S.brachypoda ,
S.kochiana Trautv., S.jenisseensis B.Floder., S.
acutufolia Willd., S.saposhnikovii A.Skvorts., S.
coesia Vill., S.rosmarinifolia L.and those species
growing in natural habitats.
The first symptoms of the disease were noted in
July.Early appearance of the disease(first ten-day pe-
riod)and active dispersion of the fungus by conidial
sporulation were recorded in the years especially
favourable to development of powdery mildew (first a
drought period and then high humidity and temperature
18—20°C).In unfavourable years the symptoms of the
disease appeared in late July.In that case the conidial
stage developed weakly , the pathogens began immedi-
ately to form the resting stage(cleistothecia).
Detailed study of the disease dynamics in 2005
made it possible to determine the difference in forma-
tion of the conidial and ascous stages.The first symp-
toms of farina were noted in the second ten-day period
of July in only S.myrtilloides and S.brachypoda.
Two ten-day periods later (2005-08-01)appearance of
anamorph was recorded in S. kochiana , S. lanata
and S.taraikensis.In S.saposhnikovii and S.di-
waricata conidial farina was formed in the second and
third ten-day periods of September(2005-09-12 , 2005-
09-30).
Later formation of the conidial stage is connected
with a higher resistance of plants to powdery mildew.
It was noted that the symptoms of disease varied
depending on a willow species.Farina of the pathogen
formed either on both leaf surfaces , or on one of them
(upper or lower).In most cases micromycetes deve-
loped mycelium on both leaf surfaces(S.coesia , S.
brachypoda , S.pyrolifolia and others).Farina formed
most intensively on the lower leaf surface.Farina on
the upper leaf surface was recorded in S. saposh-
nikovii , S.lanata and S.kochiana.
Mycelium distribution was local or as a continuous
grey-white or dirty white filmy , powdery or tomentose
farina of various density.Local or continuous farina was
also noted along the veins in willows in natural habi-
tats.Local distribution of mycelium (Fig.1)prevailed
in collection specimens , the exception was S.myr-
tilloides.In some cases growth of mycelium varied de-
pending on the leaf surface.For instance , on the upper
leaf surface of S.coesia and S.brachypoda it looked
like spots , on the lower surface like a continuous lay-
er.Disappearance of mycelium from one of the surfaces
towards the end of vegetation was noted in S.brachy-
poda , S.saposhnikovii , S.nigra and S.saxatilis.
234 菌 物 研 究 2007 年
Fig.1.Local farina on the upper leaf surface of Salix
lanata
Fruiting bodies began to form during the second
half of vegetation (second ten-day period of July)on
both leaf surfaces , chiefly on the lower one.Single
cleistothecia were found on leaves of S.saposhnikovii ,
S.caprea , S .kochiana and S.rosmarinifolia.Nu-
merous Uncinula adunca fruiting bodies developed on
leaves of S.nigra , S.myrtilloides and S.pyrolifoli-
a.In some cases(S.brachypoda)single cleistothecia
formed on the upper leaf surface and numerous on the
lower leaf surface.Fruiting bodies were not found on
leaves of S.alba , S.viminalis and some species
growing in natural habitats.Probably powdery mildew
pathogen winters as a mycelium and develops on the
shortened cycle.Predominance of the conidial stage in
the development cycle encourages larger pathogen labil-
ity and ability to utilize new host plants.
Two types of localization of cleistothecia:uniform
or in groups , often with radial arrangement , were es-
tablished.The second type prevailed.
Differences in cleistothecium formation and ripen-
ing in various willow species were observed during veg-
etation (Table 1).Collection made on August 29 ,
2005.
Table 1.Differences in formation and ripening of fruiting bodies in various willow species
Host plant Fruiting bodies
Upper leaf surface Lower leaf surface
Appendages Asci Spores
S.myrtilloides + + - - -
S.saxatilis + + - + +
S.kochiana + + + - -
S.lanata + - - - -
S.pyrolifolia + + + - -
S.rorida + + + + +
S.acutifolia + - + + +
S.daphnoides + + + + -
S.saposhnikovii + + - - -
Note:“+” presence of formed fruiting bodies , appendages , asci and spores;“ -” absence of fruiting bodies, appendages , asci and spores
In most of willow species the pathogen forms fruit-
ing bodies on both leaf surfaces , except for S.lanata
and S.acutifolia.In some species:S.saposhnikovii ,
S.myrtilloides and S.lanata cleistothecia had indif-
ferentiated contents and reduced appendages.An inter-
esting character of the pathogen was recorded in S.
saxatilis:absence of appendages , but presence of ripe
asci with ascospores.Probably the pathogen applies all
its energies to formation of wintering fungus structures
for preservation of greater deal of infection for the new
season.
U.adunca fruiting bodies ranged in size from
135 to 165μm.Appendages are numerous , equatorial ,
comparatively short (135—188 μm.), thin-walled ,
colourless , bent backwards or strombuliform at the
ends.A peculiarity of micromycete in the species of the
genus Salix from natural habitats is reduced ap-
pendages(Fig.2).Many scientists note presence of
numerous appendages(more than 120)as a character-
istic feature of the pathogen.However , our study
showed that in certain species(S.rorida , S.acutifo-
lia , S.lanata)the fungus formed a limited number of
appendages(less than 100)(Fig.3).
235 第 5卷 第 4 期 Tomoshevich M.A.:寄生于柳属植物的 Uncinula adunca 的生物学特性
a.Reduced in Salix sp.growing in natural conditions;b.Formed in the collection specimens of the genus Salix.
Fig.2.Appendages of fruiting bodies of Uncinula adunca
a.S.acutifolia;b.S.pyrolifolia
Fig.3.Uncinula adunca cleistothecium
Fig.4. Asci with ascospores of U. adunca mi-
cromycete
Asci , 4—10 in a cleistothecium , are ellipsoid ,
on short stalks , (67—75)μm×(33—45)μm , 3—
6 sporal.More often we recorded asci with 3—4
acrospores (Fig.4).Spores are ellipsoid , (20—
30)μm×(10—17)μm.
In local conditions the fungus had indifferentiated
contents of fruiting bodies in most of willow species up
to the end of vegetation.
Intraspecific resistance of willows was revealed.
For instance , out of 4 groups of S.rorida powdery
mildew was recorded in only two groups.In S.pyro-
lifolia one ecotype is susceptible to powdery mildew ,
the second one is immune.Apparently this is connect-
ed with plant age , microclimate of the site of growing
in the arboretum and origin of this group of plants.
(下转第 239页)
236 菌 物 研 究 2007 年
表 2 绣球菌在不同类型淀粉基质中的效应
Table 2. Effect of culture media with different types of
starch on Sparassis crispa growth
淀粉种类
Starch kind
生长阶段
Growth phase
原基出现率/ %
Primordium
appearance rate
ck
胶质原基略现 ,绒毛菌丝 ,菌束 0
未去皮马铃薯块
Fresh potato
pieces with peel
带皮处仅稀疏菌丝 0
鲜去皮马铃薯块
Fresh potato
pieces without peel
原基 100
马铃薯淀粉
Potato starch
胶质原基 ,菌束 50
大米淀粉
Rice starch
原基 、菌束 100
小麦淀粉
Wheat starch
原基 100
甘薯淀粉
Sweet potato
starch
胶质原基 ,菌束 50
3 讨 论
淀粉是绣球菌生长发育的重要碳源之一 ,淀
粉经绣球菌的胞外酶降解为糖类被吸收。杨瑞
长[ 4]研究表明 ,淀粉可诱导纤维素酶的活性 ,从而
促进菌丝生长和子实体形成。不同种类淀粉基质
对绣球菌生长效应不同 ,去皮新鲜马铃薯块作为
淀粉基质优于马铃薯淀粉的原因可能在于其除含
淀粉外 ,还含有微生物必需的生长因子 ,而加工淀
粉的过程中造成了营养流失。培养绣球菌如果添
加大量新鲜马铃薯块必须去皮 ,因为通过试验发
现马铃薯皮对绣球菌丝生长有抑制作用 。
据刘正南[ 2]于 1986年报道的调查资料 ,绣球
菌多生长在东北长白山 、云南等海拔约 3 000 m
的林区 ,属于山地寒湿带地域 ,平均温度 14.6 ～
12.7℃,平均降水量 228.9 ～ 148.3 mm ,平均郁闭
度 0.4 ～ 0.6。据此生存环境 ,结合试验结果可知 ,
绣球菌应属于中低温型食用菌类 ,绣球菌栽培生
育期约为 120 d ,生殖阶段时间很长 ,约占整个生
育期的 2/3 ,大体可分为菌丝生长 、原基形成 、原
基分化和子实体发育 4个阶段。绣球菌栽培管理
难度较大 ,特别是原基形成阶段处于一个特定环
境 ,即原基多出现在袋的中部和底部的微生境 ,既
需氧又嫌氧 ,称为“兼性嫌氧微生态” 。由于绣球
菌原基发育阶段存在此种特殊生理习性 ,在栽培
中必需塑造相似的生长环境 ,以保持O2/CO2 合适
的比值。
参考文献:
[ 1] 丁湖广.绣球菌生物学特性及驯化栽培技术探讨[ J] .菌物
学报 , 2005 , 24(增刊):104-106.
[ 2] 刘正南.一个珍稀食用菌———绣球菌[ J] .食用菌 , 1986
(5):6-7.
[ 3] 林衍铨.绣球菌菌丝生长营养生理研究[ J] .菌物学报 ,
2005, 24(增刊):170-171.
[ 4] 杨瑞长.米粉和石膏对香菇的增产作用[ J] .食用菌 , 1987
(5):22.
(上接第 236页)
Thus , among 60 species of the genus Salix stu-
died in the collection of the arboretum of Central
Siberian Botanical garden , SB RAS , powdery mildew
(Uncinula adunca)develops in 15 species.Plant in-
jury is of average and high degree.The disease occurs
annually and causes decrease in ornamental quality.It
has been noted that the fungus can form fruiting bod-
ies , asci and ascospores for a short period or does not
form them at all.
References:
[ 1] Yachevsky A A.Manual for Identification of Fungi.Vol.1[ M] .
St.Petersburg:Tipographia Kinda , 1913:935.
[ 2] Golovin P N.Powdery mildews parazitising in cultivated and wild
useful plants [M] .Mosсow-Leningrad:Publ.House of AS USSR ,
1960:264.
[ 3] Gelyuta V P.Flora of fungi of the Ukraine[ M] ∥Gelyuta V P.
Powdery mildews.Kiev:Naukova Dumka , 1989:256.
[ 4] Azbukina Z M.Lower plants , fungi and bryophytes of the Russian
Far East [ M] ∥Abukina Z M.Fungi.Vol. 2.Ascomycetes:
Erysiphales, Clavicipitales , Helociales.Leningrad:Nauka , 1991:
394.
239 第 5卷 第 4 期 林衍全等:绣球菌生物学特性若干问题的研究