全 文 ：第 19 卷 第 1 期
Vol. 19 No. 1
草 地 学 报
ACTA AGRESTIA SINICA
2011 年 1 月
Jan. 2011
Response of Tall Oatgrass to Salinity Stress at the Seedling Stage
YAN Hongwei1, 2 , WANG Zan2 , YANG Xi2 , WANG Xuemin2 , GAO Hongwen2*
(1. C ol lege of Pastoral Ag riculture S cience and Technology, Lanzhou University, Lanzhou, Gansu Province 730020, China ;
2. Inst itute of Animal Science, Chinese Academ y of Agricultural Science, Beijing 100193, Chin a)
Abstract: T his experiment w as conducted to evaluate the responses of tall oatgrass plants fo r salinity
st ress. T w o tall o atg rass genotypes, ZXY03P367 and ZXY03P443, w ere gr ow n fo r 14 days at g reen
house condit ions and treated w ith four N aCl concentrat ions ( 0, 65, 100, and 135 mM ) for 21 days. The
plant g row th, ion contents, photosynthetic gas exchange, lipid perox idat ion, and proline accumulat ion
w ere measur ed. M ost par ameters for the tw o genotypes w ere significant ly dif ferent af ter treated in 100 and
135 mM NaCl. Salt t reatment decr eased in roo t and shoot biomass, photosynthetic rate ( A ) and stomatal
conductance ( g s ) , and K
+
content , and increase in Na
+
content . Lar ger reduct ions in the parameter s oc
cur red in ZXY03P443. A signif icant accumulat ion of lipid perox idation and pr oline in leaves w as found
during the period o f intensive leaf grow th. These org anic compounds maybe play a ro le in leaf osmot ic ad
justment and in protect ion of membrane stability at severe salinity levels. Our results indicated that the
tw o tall oatgrass genotypes differed in their sensit ivity to salinity, with ZXY03P367 being classif ied as rel
at ively salt tolerant and ZXY03P443 as sensit iv e.
Key words: T all oatgr ass; Seedling; Salinity st ress
高燕麦草苗期对盐胁迫的响应
闫洪唯1, 2 , 王 赞2, 阳 曦2 , 王学敏2 , 高洪文2*
( 1.兰州大学草地农业科技学院, 兰州 730020; 2.中国农业科学院北京畜牧兽医研究所, 北京 100193)
摘要: 以高燕麦草( A r rhenatherum elatius L . ) ZXY03P367 和 ZXY03P443 两种基因型为材料, 研究其对不同浓度
盐胁迫的响应。幼苗生长至三叶期,分别用浓度为 0, 65, 100 和 135 mM NaCl处理 21 d, 测定 4 个盐浓度下的高
燕麦草地上、地下生物量、离子含量、光合气体交换、脂质过氧化物、脯氨酸的积累。结果表明:在 100 和 135 mM
NaCl条件下, 2种基因型高燕麦草大部分参数都存在显著差异(P < 0. 05)。盐处理导致地上和地下生物量、光合速
率( A )、气孔导度( g s )和 K + 含量降低的同时, N a+ 含量显著增加。而在叶片快速生长期, 脂质过氧化物和脯氨酸显
著积累。推测在高盐浓度胁迫下,这些有机物可能在高燕麦草叶片渗透调节和保护膜的稳定性上发挥着重要作
用。综合分析, 2 种高燕麦草基因型对盐度的敏感度不同, ZXY03P367 相对 ZXY03P443 具有较强的耐盐性。
关键词:高燕麦草; 苗期;盐胁迫
中图分类号: Q945. 78 文献标识码: A 文章编号: 10070435( 2011) 01009007
Salinity is the major environmental factor lim
it ing plant grow th and product iv ity. It also has se
r ious ef fects upon both irrigated and dr yland agri
cultural areas in China. It w as reported that up to
4. 0 107 hm 2 of land are curr ent ly affected by sa
l inity[ 1] . The development of salttolerant crops is
requisite to decr ease amounts of arable land under
going salinizat ion and to meet the g row ing human
populat ions fo od demand. T hus, w e need to iden
t ity the degree of salinity tolerance w ithin cr ops
and their w ild relat ives[ 2] . Using salttolerant fo r
age species is one method to maintaining ag ricul
tur al production in saline soils. Consequent ly, it is
necessary to identify plant species for saline areas in
both dryland and irrigated lands that w ill provide both
groundcover and agricultural production
[ 3]
.
收稿日期: 20101110;修回日期: 20101207
基金项目:现代牧草产业技术体系( 2060302) ;中国农业科学院基本科研业务费( 2011cj15) ;牧草种质资源保护项目( 2130135)资助
作者简介:闫洪唯( 1983 ) ,男,河南鹿邑人,硕士研究生,主要从事牧草种质资源研究, Email: yhwg ood@ 163. com; * 通讯作者 Author for
cor respond ence, Email : gaohongw en@ 263. net
第 1期 闫洪唯等:高燕麦草苗期对盐胁迫的响应
Salinity af fects the plants morpho logical, phy sio
logical and biochem ical processes. Salinity str ess
results in stunt ing of plants
[ 4, 5, 6]
. In such condi
t ions, shoot and roo t dr y weights w ere de
creased
[ 7]
. Zhao et al . ( 2007) r eported that salt
st ress w ith 50 mM reduced total leaf area of naked
oat ( A v ena sat iv a L. ) by 35% and plant biomass
by 52%
[ 8]
. With a higher salinity level ( 250 mM ) ,
plant grow th w as further suppr essed, causing de
creases of 91 and 86% in to tal leaf area and plant
dry mat ter , r espect ively
[ 8]
.
One mechanism associated w ith g reater toler
ance to moderately saline environments is the abili
ty o f plants to exclude Na+ f rom the shoo t, and
concurrent ly maintain high levels of shoot K + [ 9] .
Salinity can interfere with K + and Ca+ nutrition [ 10]
and cause nutr ient def iciencies. Grow ing plant
w ith NaCl st ress induces the incr ease of Na+ , and
the decrease of Ca
2+
, K
+
and M g
2+
levels in many
plants [ 7] .
Biomass product ion is a direct reflect ion of net
photosynthesis. Decreased photosynthetic r ates
may be caused by closur e o f stoma, induced by o s
motic st ress, or f rom saltinduced damage to the
photosynthet ic apparatus
[ 11]
. Wilson et al . ( 2006)
reported a highly significant reduct ion in stomatal
conductance ( g s ) , and net pho to synthet ic r ate ( A )
in cow pea ( Vigna ung uiculata ( L. ) Walp. ) cult i
var s suffer ing from salinity. M oreover, the net
photosynthet ic rate w as a more sensit ive phy sio
logical indicator of the level of salinity st ress than
the leaf area
[ 12]
. In rice ( Or y z a sat iv a L. ) , A was
decreased by about 35% at 12 ds m- 1 w ith salt
st ress, but the reduct ions in g s and tr anspirat ion
rate ( T r ) w ere 74 and 63%, respect iv ely . Con
versely, the reduct ion in intercellular CO 2 concen
t rat ion ( C i ) w as much low er ( 15%) at the same
st ress level[ 13] .
Tall oatgrass, native to Europe, is cult iv ated
in the gr asslands of the central and nor thern USA,
w estern Eurasia, East Asia, and the M editerra
nean basin[ 14] . T al l oatgr ass is a useful conserva
t ion g rass for cover and forag e on surface m ined
lands and mar ginal pastureland, and can also be
used for livestock fo rag e beg inning in its second
g row ing season. Because of it s considerable r esist
ance to drought , cold, disease and insects, tall
oatgr ass w as o ften used as a tert iar y germplasm re
source for improving oat w hich is the most closely
related genera to A r rhenather um w ithin the tr ibe
Aveneae
[ 15]
. The Poaceae family ( grasses ) is
known to include genera that range f rom moderate
ly salt sensitive to highly salt tolerant [ 16] . L it t le in
format ion is available on the responses of tall oat
gr ass to salinity[ 17] . Understanding the mecha
nisms under lying the g row th and physio logical re
sponses to salinity w ould be valuable in select ing
and improv ing tall oatgr ass str ains tolerant o r a
dapted to salt st ress. Our present w ork w as de
signed to determ ine the inf luence of salinity st ress
on seedling gr ow th, ion content , photosynthetic
productiv ity and metabo lism of tall o atg rass.
1 Materials and methods
1. 1 Plant Materials and Salt Treatments
T w o tall oatgr ass geno types, ZXY03P367
and ZXY03P443, w ere selected fo r this study. A
2 4 factor ial experiment , arr anged in a complete
ly random ized design w ith three r eplicat ions, w as
conducted. Tw enty seeds w er e planted in twoliter
pots f illed w ith silica sand and thinned to ten
plants per pot af ter emergence. Pots w ere irrigated
w ith dist illed w ater for 7 days and then fert ilized
w ith a saltfree Hoag land s so lut ion. Salinity
str ess w as impo sed on seedling s at 14 days of age
by adding 65, 100, and 135 mM N aCl in saltfree
Hoag land s so lut ion, w hile taking the saltfree
Hoag land s solution itself as the control t reat
ment. For each pot , 200 mL o f salt solut ion w as
applied daily to ensure that all seedlings receiv ed
an equal volume of t reatment solut ion and to pre
vent addit ional drought str ess. T he exper iment
w as performed in a greenhouse w ith a temperature
of 25/ 16 ( day / night ) and a 16h photoperiod
w ith 300 mo l m- 2 s- 1 illuminat ion. T he rela
t ive humidity was maintained at about 70% . The
plants w ere then subjected to the salt t reatments
for 21 day s before the fol low ing measurements
w ere taken.
91
草 地 学 报 第 19卷
1. 2 Estimation of Plant Growth
Plant shoots and roo ts w er e harvested af ter 21
days of salinity t reatment, and dr ied at 60 for 48
h fo r the determinat ion o f dry biomass and further
analysis.
1. 3 Estimation of Ion Content
Dried leaf samples w er e ground to a f ine pow
der and approximately 0. 1 g w as t ransferred to a
test tube. Ions w er e ex t racted by adding 10 mL o f
0. 1 N acet ic acid and heat ing in a w ater bath at
80 for 2 h. The ex t ract ion solut ion w as coo led at
room temperatur e and left overnight, and then fil
tered using No. 40 Whatman f ilter paper. Sodium
and potassium concentrat ions w ere then deter
mined using an atom ic absorption spect rometer
( Perkins Elmer, Norw alk, CT, USA) .
1. 4 Estimation of Photosynthetic Parameters
Net photosynthetic rate ( A ) , and stomatal
conductance ( g s ) w ere measured on the same leaf
samples as the leaf greenness measur ements using
an inf rared, open gas exchange sy stem LI6400
( LICOR) fo llow ing the manufacturer s inst ruc
t ions. T he area of each leaf in the pho to synthet ic
meter chamber w as determined manually. T he
measurements w er e perfo rmed w ith light levels o f
( 1000 mol m- 2 s- 1 ) . Data w ere manually re
corded when the gas exchange parameters became
stable.
1. 5 Estimation of Proline and Lipid Peroxidation
Free pr oline w as quantif ied spectr ophotomet
r ically using the method o f Bates et al .
[ 18]
. L ipid
perox idat ion w as est imated by measuring the con
tent o f 2thiobarbitur ic acidreact ive substances in
0. 2 g leaf f resh w eight acco rding to M adhava Rao
and Sresty[ 19] . M alondialdehyde ( MDA ) content
w as determined spectrophotometrically at A 532 and cor
rected for nonspecif ic turbidity at A 600 and A450 .
1. 6 Statistical Analysis
All data w ere subjected to analysis of v ariance
using the general linear model pro cedure of SAS
( SA S, 1996) . T reatment mean dif ferences w ere
separated by the least signif icant difference ( LSD0. 05 )
test if F tests w ere significant ( P 0. 05) .
2 Results
2. 1 Plant Growth Response to Salinity
Shoot biomass of tw o tall o atg rass genotypes
w as signif icant ly decreased with increased NaCl
concentrat ion ( Table 1) . Average of 29% r educ
t ion in shoo t biomass w as found at 65 mM salinity
level compared to the contr ol, and fur ther de
cr eased w ith higher N aCl concentrat ion. In 65 ~
135 mM NaCl st ress, decrease in shoot biomass
ranged fr om 28% to 71% and 30% to 80% fo r
ZXY03P367 and ZXY03P443, respect ively. Mo
reover, signif icant dif ferences in aboveg round bio
mass production w er e observ ed, bo th betw een gen
otypes ( P< 0. 05) and salinity st ress ( P< 0. 001) .
No obvious genotype salinity interact ion meas
urements w ere found.
Root biomass of tw o tall oatg rass g enotype
exhibited a similar t rend to shoo t biomass w ith salt
str ess ( T able 1) . On average, a 30%, 65%, and
86% reduction in root biomass occurred in 65,
100, and 135 mM salinity lev els, respectively,
compared w ith the control.
T able 1 Shoo t and ro ot biomass o f two tall oatg rass genot ypes under four salt t reatments measured at the seedling st ages
Material s
S hoot biomass
0 mM 65 mM 100 mM 135 mM
Root biomass
0 mM 65 mM 100 mM 135 mM
ZXY03P367 1. 96 0. 03 1. 42 0. 03 1. 02 0. 06 0. 56 0. 05 1. 25 0. 02 0. 87 0. 05 0. 43 0. 04 0. 20 0. 04
ZXY03P443 1. 52 0. 04 1. 07 0. 09 0. 73 0. 07 0. 38 0. 08 1. 00 0. 02 0. 70 0. 04 0. 34 0. 05 0. 12 0. 02
Genotyp e ( G) * *
S alin ity ( S) * * * * * *
G S ns ns
Note: * and * * * mean signif icant dif feren ces at the 0. 05 and 0. 001 probabilit y level s, r espect ively. ns means nonsignif icant dif f erence at
th e 0. 05 level
92
第 1期 闫洪唯等:高燕麦草苗期对盐胁迫的响应
2. 2 Sodium and Potassium Content
With salinity st ress, leaf Na
+
contents in
creased significant ly ( P< 0. 001) w ith increasing
NaCl concentrat ions ( F ig . 1A) . H igh Na+ accumu
lat ion was observ ed in leaves of ZXY03P367 and
ZXY03P443. Compared w ith the control, Na +
accumulat ion in ZXY03P443 increased by about
3. 2, 15, and 24. 7 t imes in 65, 100, and 135 mM
NaCl st ress, respectively, w hile these values w er e
2. 7, 13, and 23 times for ZXY03P367. M oreo
ver, in ZXY03P443, the increase in Na+ content
w as compar at ively higher than ZXY03P367 in all
salt t reatments ( F ig. 1A) .
Increasing salt levels led to a signif icant r educ
t ion in leaf K + concentrat ion ( F ig. 1B) . T he r educ
t ion o f K + in ZXY03P443 w as g reater than
ZXY03P367 at all salt levels ( F ig. 1B) . As a re
sult , leaf K contents w ere slight ly higher in
ZXY03P367 than in ZXY03P443. How ever, no
dif ference between the tw o species w as found in
any salt t reatments.
Fig . 1 Effect s o f increasing NaCl concent ration on shoo t Na+ ( A) , K + ( B) , pho tosynthetic r ate ( C) , stomatal
conductance ( D) , P roline ( E) , and MDA ( F) content of two tall oatg rass g enotypes, ZXY03P367 ( open columns) and ZXY03P443
( clo sed solumns) , exposed to four NaCl t reatments o ver 21 days. Ver tical bar s represent S. E. of t reatment ( n= 3)
2. 3 Photosynthetic Gas Exchange
Leaf photosynthet ic rate ( A ) and stomatal
conductance ( g s ) were significant ly higher in
ZXY03P367 than in ZXY03P443 ( Fig . 1C, D) .
With sal inity st ress, leaf pho to synthet ic rate w as
reduced significant ly ( P< 0. 01) fo llow ing the in
creased NaCl concentrat ion ( Fig. 1C) . Signif icant
reduct ion in A was observed even at the low salini
ty levels ( F ig . 1C ) . The A decr eased by about
21% , 57% , 76% when seedlings w ere subjected to
salt st ress of 65, 100, and 135 mM , respect ively;
how ever, reduct ions in g s w ere slight ly low er than
in A , amount ing to 17%, 31% and 61 % w ith the
same salt levels, respect ively ( F ig. 1D) .
2. 4 Accumulation of Proline and MDA
Pro line concentration w as measured at the
seedling stag e to determ ine its associat ion w ith sa
l inity tolerance. With salt st ress, proline concen
t rat ion increased signif icant ly in the tw o genotypes
( F ig. 1E) . On average, a 83% , 244% , and 262%
increase in proline content w as found at 65, 100,
and 135 mM salinity lev el, respect ively, compar ed
to the contr ol.
NaCl t reatments signif icant ly increased MDA
content in both tall oatgrass genotypes, w ith the
higher relat ive incr ease observed in ZXY03P443,
in w hich it increased 18. 1% ~ 97. 2% , w hereas a
much low er increase w as observed in ZXY03P367
( 6. 3% ~ 48. 4% ) ( Fig. 1F) . It indicated a higher
level o f lipid pero xidat ion in ZXY03P443 due to
salt st ress.
3 Discussion and conclusion
Shoot and roo t gr ow th inhibit ion is a common
response to salinity, and plant g row th is one of the
most impo rtant agr icultural indices of salt st ress
93
草 地 学 报 第 19卷
to ler ance as indicated by different studies [ 20, 21] .
Salt st ress signif icant ly reduced the grow th of the
tw o tall o atg rass genotypes during the seedling
stage. T he observed reduction in shoo t and root
biomass is likely to be due to a combinat ion of slo
wer grow th and development as a result of osmot ic
st ress [ 22] and an inhibit ion o f photosynthesis either
as a result of the direct ef fects o f salinity on the
photosynthet ic apparatus or the indirect ef fects of a
reduct ion in sink capacity
[ 23]
. ZXY03P443 w as
found to be mo re sensitive than ZXY03P367 to sa
l inity t reatments. H ow ever, the dif ferent ial re
sponse of g row th to salinity could be due to g eno
ty pic differences[ 24] . Genotypic dif ferences to salin
ity have been reported in gr een g ram[ 25] , bar
ley[ 26] , r ice[ 13] .
The reduct ion in photosynthesis w as small
w hen plants w ere subjected to salt levels low er
than 100 mM ( Fig. 1E, 1F) , but significant effects
occur red at higher salt concentrations. This is sim
ilar to Zhao et al . ( 2007) findings for rice. T he
change in g s was sim ilar to that o f A in tal l o at
g rass[ 8] . A close relat ionship w as found betw een A
and shoo t biomass and g s ( F ig . 2) , suggest ing that
the sever e reduct ion in g row th w ith salt st ress w as
st rong ly related to the reduction in leaf gas ex
change propert ies. Analysis of g as exchange meas
urements revealed that the greater relat iv e reduc
t ions in A than in g s ( F ig. 1C, 1D) measured in tall
oatg rass suggests that nonstomatal inhibition o f
photosynthesis, caused by direct ef fects of NaCl on
the photosynthet ic appar atus independent of stom
atal closur e, m ight be responsible for the r educt ion
in photosynthet ic r ate. Nonstomatal inhibition o f
photosynthesis by salinity w as also reported for
several other plant species
[ 12]
. H ow ever, other re
searchers indicated that photosynthesis w asnt de
creased by salinity and even stimulated by low salt
concentrat ions
[ 27]
. In A . p seudoalhagi , the leaf
CO 2 assimilat ion rate increased in condit ions of low
salinity ( 50 mM NaCl) and is not signif icantly af
fected by 100 mM NaCl. It is , how ever, reduced
to about 60% of the contro l by 200 mM NaCl. T he
observed decrease in both g s and tr anspiration rate
m ight be among the impo rtant adapt iv e mecha
nisms conferring to lerance to salinity in rice
[ 13, 28]
.
Decreases in the photosynthet ic rate are due to sev
eral facto rs: ( 1) the dehydrat ion of cell membranes
w hich reduce their permeability to CO 2 , ( 2) salt
tox icity, ( 3) the reduction o f CO 2 supply because
of the hydro act iv e closure of stomata, ( 4 ) en
hanced senescence induced by salinity , ( 5) changes
in enzyme activity induced by alterat ions in cy to
plasmic st ructure, ( 6) negative feedback by re
duced sink act ivity[ 29] .
Fig. 2 Relationship betw een photo synthet ic rate, and
stomatal conductance ( ) and shoot biomass ( ) in tall
oatg r ass. ** and *** significant differences at the 0. 01
and 0. 001 probability levels, r espectively
An inter est ing characterist ic of most plants
g row ing in saline environments is the accumulat ion
of proline
[ 30]
. Our results show ed a substant ial in
cr ease in pr oline concentrat ion w ith less than 100
mM N aCl t reatment. A significant ly high accumu
lat ion of proline was probably associated w ith os
motic adjustment and protect ion o f membrane sta
bility[ 31] . How ever, Lut ts et al . ( 1996) observ ed
that proline did no t show relat ionship to an o smo tic
adjustment in salt str essed rice plants[ 32] . In some
studies, the accumulat ion of proline has been relat
ed to the ionic status o f the plants
[ 33, 34]
. Colmer et
al . ( 1996) show ed that high proline content w as
associated w ith the maintenance of a more favoura
ble K + to Na+ ratio in the NaCltr eated ro ot t ips
supplied w ith Ca2+ [ 35] . In this study , a high posi
t ive corr elat ion w as found betw een proline and
Na+ concentr at ion in leaves ( Fig . 3) , suggest ing
94
第 1期 闫洪唯等:高燕麦草苗期对盐胁迫的响应
the positive role of Na+ in pro line accumulation.
The same r esult w as also observed in S esuvium
por tulacast rum
[ 36] . The contribut ion of the proline
to the osmot ic adjustment becomes quite signif icant
by the fact that this compatible osmolyte is concen
t rated most ly in the cytosol and the chloro
plasts[ 37] .
Perox idat ion of membrane lipids is an indica
t ion of membrane damage and leakage w ith salt
st ress condit ions
[ 38]
. Salt st ress af fected both gen
o types by means of lipid perox idation but ,
ZXY03P443 had higher incremental rates o f all
NaCl t reatments. Grow th inhibit ion w ith salinity
in ZXY03P443 is in good co rrelat ion w ith in
creased l ipid pero xidat ion lev els. Low er MDA level
w as remarkable in ZXY03P367 even at the highest
NaCl concentrat ions ( F ig . 1D) . It is presumed that
the ex tent of membrane damage is not so severe in
ZXY03P367 due to salinity. This is consistent
w ith the results obtained for Na
+
and K
+
content .
Sim ilar results occurred in salt to ler ant barley cul
t ivars [ 39] , and salt resistant tobacco plants also had
low er lev els o f lipid perox idation w hich is an im
portant sign of higher ox idative damage lim it ing ca
pacity w ith salinity[ 40] . How ever, salt sensit iv e
rice variet ies had higher MDA content and elect ro
lyte leakage in response to salt st ress
[ 13]
. In addi
t ion, a high positive co rrelat ion w as established
betw een MDA and Na+ concentrat ion in leaves
( Fig . 3) , suggest ing the po sit iv e role of Na+ in
MDA accumulat ion.
Fig . 3 Relationship betw een Na+ content, and Pro line ( ) ,
and MDA ( ) content in tall oatgrass. ** and *** significant at
the 0. 01 and 0. 001 probabilit y lev els, r espectiv ely
In conclusion, salinity st ress signif icant ly in
hibited the grow th of tall oatgrass. Betw een the
tw o geno types in their responses to salinity , there
are signif icant differences w hich ar e consistent
w ith their physio logical responses measured in dif
ferent salinity levels. T he general g row th, g as ex
change, lipid pero xidat ion and pro line accumula
t ion of tw o tall oatg rass genotypes are also in good
correlat ion w ith each other, indicat ing that
ZXY03P443 is mor e sensitive to salinity than
ZXY03P367.
Acknowledgments
T he authors g ratefully thank Beihong Wang
for ex cellent technical assistance and Dr. M ar shall
Haferkamp fo r his useful comments on an earlier
version of this manuscript .
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