全 文 :Characteristics of Seed Dormancy, Dormancy
Breaking and endi in at ion in Halophytes
ZHAO Ke-Fu FENG Li-Tian FAN Hai
( ~ ~ t i r u l e o f P l M f S t r e s s , ~ N o r m a l Uniwsi%y, ~inanwx)14)
Abstract The characteristics of seed dormancy, dormancy b~aking and gemination in halophytes
are discussed. The main contents are as follows : ( 1 )The seed dormancy types of halophytes ; (2) The
essential conditions for dormancy breaking of seeds in halophytes ; ( 3 ) The correlation between seed
gemination of halophytes and salinity. The inhibiting machanism of halophytic seeds by salinity is
also discussed. Lastly, several key ~roblems remaining to be solved in future researches into seed
germination of halophytes, as well as the sidcance of the study on seed dormancy and germination
of halophytes, are sumrnarised .
Key words Seeds of halophytes, Seed dormancy types, Domcy-breaking conditions, Salinity and
gemination
*%@f%%Ei%$fi@~&f % B ,%B 396783,
a &%S A, Author for w ~ e n c e .
&%6 S )gJ : 199843-30 &2 El #I :199807-29 Bff %% : X%E
1 a&s##9m#E%a
F@ 1%9 +,Nikolacra ~%@#!JR%S&%%#!lH~m#BE%B!, @GX%--!!%@
B%fiik&%%BRT#flR% 4 #%a, lVWiT!#UR.BBR(EGE
73) \%&*B#B@@%B#jR(3Z%#BE) ,S*L%M4F&*&%#3=0
&B#lRHHFWGaBBHB%%3)2BmE,iR@2E@m*E%*SBEE@
~ ~ ( $ u ~ t ~ j ~ ; n / ~ ~ ~ ) w a ; t n f i m ~ . H T ~ & B # B R & ~ + * ~ , ~PIFRBBE.++#
flR+U%#flRo -~~F%#tlRM&&~%HT%H-&RC4~~EW~B~P~UQRGii7~
HE,@--&+%#RbB#BRN&&%@HF, A % ~ F J R @ E W ~ ~ @ ~ @ @ % G Q ~ ~ J ~ #
BEo SE&B#lEMHFH#N%RG,Em&E%%A, FP@E?&&EiBlE, e&&@
QHB,%E&Eo
H & @ # u E H F m & % % @ & z % * & H T M d % N + 0 @ I # * %?*I4
(Ai~oaceae ). ?.3 $I ( Amamnthaceae ) .% 3 % ( Brassicaceae ) .6 .fSI $I ( Caryophyllaceae ) . %
(~heno@aceae),$@% (Cyperaceae). A ~ R ( E u p h o I f , i e c ~ ) . i F 7 t i ~ ( P o a c e a e ) %Z&$I
(Scmphuhiaceae)o 3&@BS~&E~&%.&%@.@~.hB@%@&%Eo
&&@@irrBBR#3@B2;t~XEEB;I;fE3),SEG@&E3J---X&f ,+P
em^^ ~ ~ E ~ E B I - Z ~ Y G ~ H ~ E ~ ~ B B R ~ ~ ~ H F E R E ~ ~ ~ ~ H
3 E ~ - 2 m * % ~ B ~ ~ N ~ \ @ M ~ ~ & ~ ~ T * ~ i l 1 m ~ ~ u % ~ * E o
SH%~M%&@~HFSUT~;~BH&@+BG&,@J~U,~Q%(~I~~-
daceae) .$B%(~~iaceae) .X$fl%#(Maceae).@?#(Arecaceae), @&&%#-(Hydrophyl-
laceae) .Hea;C(C(Liliaceae) M%Z% (Papaveraceae)
&&~~+mBz+a~BEHFs-~~zfiu~a*~#n~~e%amHT, BPS
H H F ~ E ~ @ E B ~ % % , ~ ~ R ~ ~ ~ I W E ~ ~ E I : & ~ # E M ,G#Jfi%ig33BjT;n/%
?$h9RRJk@~%%Eo #%B@#%iq%@(1985)~$A%@(~~artina angl ica)HF%
gfrtJR%,E%22SiXZ%MHFE%$SB ,E3L@Ak#FB3$k;t,E3L@/J\m#
FE~$E&2*EEB5BmHT%$E~%,B*sMTBz&@#BEmHFo
fi~H(ArnaryIlidaceae) .$B%(~~iaceae).X$~$I(Araliaceae).%#&(Arecaceae).
B @ $ I ( ~ ~ d r o p h ~ h ) .a&N(~iliaceae)%@ge&a;C(Pa~averaceae)~N%%&&&@NH
FJSlT3%B&&B#flRfreJHFa
&&&@+mS%#R(%B#N)H32fiU$E%E9SZRS%H%&EN ,S
~ ! . ? i l ~ + 3 % 5 1 3 ~ t l g t r 4 ~ 9 ~ ~ ~ ? # ~ &3S%4n8%HF&%--E;l;B7k&HtS@fE%E
BmE,E7ti%s6#URm9&Zs~TH&BkHF@7k%&5%Z,4~RSHHF%
Sk%% EI ~ ~ ~ ~ % ~ & R H B & ~ ) ~ # $ I , @ # T B ~ ~ ~ ~ J ~ ~ % ~ I ] ~ ~ ~ ~ ~ P ~ % E ~
bk%fPst%RHT%Efi&~%~bk~84F,B~BB.&~,%%i*,4~%S%84F;f;fi&MW,
HF&5%%Eo &X5&&%@+&GS-9%G@mHF9EP&G627kmH&jT;n
*BLm#nEo
@@$@ (~nacardiaceae) ,* #I % ( Bombacaceae) . J% % ( Convolvuaceae) . #8 %$ $! ( Mal-
vaceae) . ~ , % F % ( ~ a ~ i n d a c e a e ) ~ # l # % ( ~ i l i a c e a e ) 9 !HS!%&!i%FTNF&%R#IR KJH
BBB3%7k+~G~$~~~RB-~%,3HM$J#HH@%,~JUB%N%&&@B
H3&m%JHaH!BBB&EB&H(% 6 ) o
0.0% NaCl 0.5% NaCl
1.0% NaCl
1001
2.0% NaCl
1.5% NaCl
"1
%#A&&%@w&,m#@3aa2+5w&mB&n%%? &BM$J&&@@
H ~ a z H a a ~ + z ~ + ~ ~ ~ , A ~ ~ G u ~ n ~ ~ a :
3.1 EFJb!%
9% 1859% Darwin BPSsBH3BG5wZM%;tfcHf3&fr,Itf;G,~arshne~ jMI Baijal
( I ~ ~ ~ M ~ G R T B W T ? ; E & + , B R & X ~ ~ I I H ~ E B G ? ; w m & T o a , s e r w
H % l l ! % k f b J ! % ~ ~ : ~ a ~ C Q > Na2CQ> NaCl> CaC12, S~ov(l964;1974)&%&-E~-
a & T x ~ & & % % ~ l k B % ~ & ~ a o
8 4 N ~ ~ H ~ ~ ~ # Y M W ~ ~ @ I ~ ~ M R % R B ( % ) 4b-9# &H%?&*4F&&&
N ~ c ~ % B ( % ) @b!J %#%Go Uhvits (1946) I% Redrnann
fl%
0.0 1.0 3.0 5.0 (1974)H%&Hg%$ ( ~ e d i c a ~ o g i v a ) # T
%-3 f i & S i % a , 2 S i NaClW%#&33--&&
Myers %I ~ o r ~ a n ( 1 9 8 9 ) ) t i t ~ , # & l % LlpZuchnefiLFca & l%NaCl +,##T%2$hk 70%@
31 5O%,fiU%& 2 % N a C l % # T J ~ E i & , ~ U ~ ~ ~ 7 % o %@W~i%B$jHI ~ a ~ l ~ ! k ~ & l %
%%,EU~fII~%#MH3%2W%GBBB%Gm;l;R%-RT%%o Ungar(1971)
iFJRGthNXthBBPJ4kW S d depressa N ~ S ( h c i n e U i u ~ l i a n a ) , f i ~ i 3 % ( ~
7)&~sB%~%Ir$I&WEHE@BB$~;l;R~-RTW#Ro
3.3 xema
El 3 @S(S.Maritima var. ~ a c 1 w a r p a ) ~ ? @ 7 k ~ % . f f ~ B ~ . f f ~ ~ ~ ~ (UW j#I Boucaud,l975)
A. $$TMBfi#F;B. B%@*+dtF3%BfiHFiC. B 0.85 moVL NaCl +dtAXBCJHF.
macmcurja, S . rnaritima var.&ili $US. depmsa fttl@3R 0.85 moVL NaCl !dSE, h@%l
i%3XM%&%&o &ABB,B3SqU%@1%&BTSX$,MEB8BEjMr/J\%HF
BE0 %4%%%% Ungar $U Binet(l975)$ii!iBM&!k@% Spergularia media, Salicania eu-
mpaea,Suaeda sp.@%%?Bo Boucad Ungar(1976),Ungar(1977a)jMr Ungar(1978a)#f$g
i%3J%;f;flB@3%2,@ GA3 &$.fiil1B%TBqUBB,3Itl;%~~%~f~Qfi&~~
ua \B%\
1 62.22 56.52 37.78
BK4%%%!ka%%Jt!kjMrE@~B!M@a
2 48.89 54.76 37.78 RElils$2%M, BtGMFat -&T 4 .3 EB
3
8%
60.00 51.11 18.75 ?&bBq%E 9 %% ,gq%dh%gg@%X$
4 55.81 52.17 42.86 , F & 9 @ , Prisco$UVicira( l976) ,~ .~&Xf
5 60.00 54.35 35.42
57.76 53.78 34.52 HFiEE3EGM3JHR&T7k#F-%6%
1 59.52 40.91 28.07 MltJPPfbJfi;f;SEBbBdh%M#%1 ,EB! 0.1
2 53.33 46.67 29.17 m o l / L N a C l & B ; f ; $ E ~ b J # h i E ~ b % f i ,
3 63.64 45.65 30.95
9 % 4 50.00 54.55 27.66 8P@~HB~~2~)rp%1#3bHBb%B0
5 62.22 46.95 21.28 a l t f ; , ~ ~ f j m m a E s a & s r S u B k
T 57.74 46.95 27.43 N a C l ~ $ r f ~ % / ! i f F ~ , , # 3 b ~ ~ a R q
% e ~ ~ A % P ! k m & B % o %*$
&%@BiFndF&!k!/&kB%EM#&B2fl , % ~ I ~ ~ & M E E J L F B & B % I ] , A@
@ flowers(1972a) reen en way $U Osmond(1972) UW Hall %(1974)%% , & ! k @ % & f i @ ~
% 5 3 g % B & B % M , & % q % E G T % % + , % e - & ~ J ~ s ~ m @ * & - & ~ 8 ~ #
?&7 &THSS%&TaSHF%Z@(%) % 3 b , % ~ @ % H F % E & 4 3 % % ,5
SSR( - E ~2%BBMk%@B#hBa@#Bft
4 6 12 16 25 && % , Steward jMr Lee ( 1974) u 2 flowers
Nu!z% 83.6 79.2 40.0 1.6 o (1972b)%$%W$RqMBfj@B%8&%
NaHC4 88.0 87.2 31.0 2.8 0 E G % % % q i $ s & i % # H , p % @ % %
NaCl 90.0 78.8 34.0 5.2 0 To Barmnum jMr Polijalcoff-Mayber( 1977) 2 3%
%LIE$ 77.4 77.2 30.0 2.0 0
&%W;rkrfrBCldEl% 86%(91 Q Macke N Ungar,1971) 1ooumoVL !#JRi%?qU@@ 120 umoVL NaCl
fi%@$U%&$C@qU@8 12 urnoVL NaCl q@T@J%20
RLY~~~S,%&%@%HF$~~B#&G~RSB~%% , $ ? ! $ ~ H F M L B ~ B , S
%BUB&@!k&B!$P&~A#FM#Ro
%T&&&%HF#lE,ERHF#BE#%EHR% ,S$@% ,R#4E%iB ,&&&
@HF%EHB%%%B*,mBH%%E#rB&tE5@,g%EBT%3mH-e%
fi9@flG&i+$rd@$%B,@J3U&BBM&g&GE&TGHf%7k%,5B%-HR
FWt? ES%Y%H@%,5@@Z~~~~rl@%? E& TAP%&%d&HPP%1,i5E
&% DNA $$3& RNA @%@J#%j? 5E%!!!?~F&J~f?fi~? %4%#E%f%%3%k!k
&%HFREHs,&ldao
~&~H~~&,x$B-$T~&&&~HF#BE#REH$E,&&~~HE&~
@,UBF*KE$lH&&&@#8G8%&Xo
HB1 ,I+AE, 1985. Z?iJZA++#I,X@WResrlBb-22 +%esrlW%B%ikft% ,159- 170
WiEB, VBnR, &8$;3,1985. K%A++R,XPW%&iEE-22 F%&W%ieRRB*B ,171 - 178
B&er R S , UngarIV,1989. Gu,JBot,67:1420-1425
Beskin C C, B&in J M, 1995. Dormancy t y p and dormancy-breakmg and germination requirements in seeds of halophytes. In "Bi-
ology of Salt-Tolerant Plants" ( MA Khan and IA Ungar eds) . Chelsea, Michigan, USA, pp .23 - 30
Boucaud J, Ungar I A, 1976. Phpiol P h , 37 : 143 - 148
B d e S W, 1995. How do halophytes o v m m e &ty? In "Bidcg of Salt-Tolerant Plants" (MA Khan and IA Ungar eds) .
Chelsea, Michigan, USA. pp. 199 - 213
ChoudhuriGN,1968. WahhxaEodogy,49:465-471
Damin C,1859. On the origin of species by rmm of natural Selection. Jy Murray. London.
F l m T J , 1 9 7 2 . J&pBot,23:310-321
Flowem T J , 1972b. Ph*, 11: 1881 - 1886
Greenway H, Osmond C B, 1972. P h Physwl, 49 : 256 - 259
Hadas A,1976. J&pBot,27:480-489
H a l l J L , Y m A R , F l m T J . 1 9 7 4 . p f ~ p i o l , 7 1 : 2 0 0 - u ) 6
XhanAA,1971. Sciena,171:853-859
Nikolaeva M G, 1%9. Izdaterstvo "Naaka" , Leninpd (Translated from Russian by Shapim Z, National Science Foundation, Wa&-
ington, DC (USA) .
PrisooJT,VichGHI,1976. P h p i d P h , 3 6 : 3 1 7 - 3 2 0
RisooJI ,SwtoGE, Fe~eiraLGR,1978. PhntSoil,49:199-206
R e h R I . 1 9 7 4 . CMJBot,52:803-808
Steward GR,Lee J A,1974. Planta,l20:279-289
Stmpov B P. 1964. Physiological back of salt tolerance of plants. IPST J d e m .
S-v B P, 1974. Structure and function of plant ells in saline habitats John W i d sons, New York.
UngarIA,1971. Rhodora,73:548-551
Ungar I A,Binet P,1975. Aquatic Bot,1:45-55
Ungar I A, Boucaud J , 1975. Action des f a tencurs en NeCl sur revolution des c y t o h e a au- de la g e m h i o n d urn halophyte
IC Suaeda maritima (L) Durn var macrocarpa Moq C R. Acad Sci Pa&, 281: 1239 - 1242
UngarI A,1977. AquaficBot,3:329-335
UngarI A,1978. BotRev,44:233-264
Ungar I A. 1%. Seed Germination and Seed-bank in halophytes. In "Seed Development and Germiantion" (Jaime Kigel and Ged
W eds) . pp 599 - 628
V ~ K A , B a i j a l B D , 1 9 7 7 . 6mpPhysidEcdogy,2:104-107
Zhao Kefu , Feng Litian, Fan Hai , Zhang Shengqiang, 1997. Study on the seed g&on and its basic physiology in habphytm.
Supplement to h e J d of Sun Y a m University. 17