全 文 ：第 17 卷
第 1 期

Vol. 17
No . 1
草
地
学
报
ACTA AGRESTIA SINICA



2009 年
1 月

Jan.
2009

博士论文摘要

种子发芽生态研究
博士生:张红香1
导师:周道玮2
( 1.中国科学院植物研究所植被数量生态学重点实验室, 北京
100093;
2.中国科学院东北地理与农业生态研究所, 长春
130020)


生活史策略是植物种群生态研究的重要领域,
而种子发芽是植物生活史的重要阶段, 因此, 研究种
子发芽规律及对策,有助于认识和阐明物种进化及
其生态适应特征。本论文以松嫩草原区植物为对
象,主要研究种子对恒温和变温的发芽响应, 比较变
温梯度模拟的全球增温对 C3、C4 植物种子发芽的
影响,研究植物发芽模式及在不同温度条件下的变
化,探讨植物对盐、水势变化及盐与温度交互作用的
发芽响应机制。
根据恒温对种子发芽影响的结果, 在积温模型
的基础上提出了修正积温模型 ( modif ied thermal
time model) , 补充了发芽速率恒定部分的拟合公
式。C3 植物的发芽速率明显低于 C4 植物。根据积
温模型预测的发芽最低温和积温常数, C3 植物的平
均发芽最低温低于 C4 植物,平均积温需要明显高于
C4 植物。本研究指出, 种子发芽类型可以分为五
类:速萌型、延迟型、稳定型、正态型、延迟速萌型。
多数一年生植物属速萌型, 分布于草甸草原的多年
生植物属延迟速萌型、正态型或稳定型。大部分植
物在不同温度下的发芽类型变化,是对环境的一种
适应对策。
在模拟全球增温的变温梯度条件下, C3、C4 植
物的发芽特点均发生改变。C3 植物对高温敏感, C4
植物对低温敏感。根据几个变温梯度的比较, 全球
增温不是温差变化,而是高温或低温部分的温度影
响种子发芽。野外条件下, 松嫩草原区更多的植物
种子在春季而非秋季发芽。室内实验证明,在春、秋
季温度范围( 5~ 10
和 10~ 15
)内, 日温递增时
发芽显著高于递减时, 植物春季发芽是自然选择的
一种适应策略。
虎尾草( Chlori s v ir gata Sw artz) 和马唐( Dig-
i taria sanguinal is )在不同温度下发芽速率和盐度
之间成曲线关系, 认为可能存在生理机制,使得中间
盐度和高盐浓度下的渗透胁迫降低, 发芽速率明显
增加,产生了曲线而非直线的关系。进一步实验表
明, PEG和 N aCl两种水势处理相比,两个物种在更
低的 NaCl引起的水势下发芽, 且比在等渗的 PEG
中快。实验用植物种子的干重随时间降低, NaCl处
理中发芽种子的含水量、钠离子浓度和种子溶液钠
浓度显著高于 PEG处理。NaCl处理中发芽种子和
未发芽种子钠离子浓度, 种子溶液钠浓度均随水势
降低而增加。种子含水量、钠离子浓度、种子溶液钠
浓度随时间变化呈线性增加。
根据实验结果提出盐度模型:
S = ( S m- S ) t ,其
中
S 是盐度常数, Sm 是种子能够发芽的最高盐度
值, S 是外界盐浓度, t是某个盐浓度下的发芽时间。
将种子对盐浓度的响应划分为四个阶段:第
阶段
主要是渗透效应起负作用; 第
阶段是渗透效应和
离子效应共同作用, 离子效用的正作用强于渗透效
应的负作用; 第
阶段等渗的 PEG 中已无种子发
芽,离子效应的正作用远远高于渗透效应的负作用;
第
阶段是离子效应开始起到离子毒害的负作用。
本论文结果对于预测全球变暖对植物生长分布
和发芽出土的影响, 开发利用野生植物资源策略均
有理论指导意义和生产实践价值。
关键词: 种子; 发芽;温度; 积温模型;盐; 渗透效应;离子效应
中图分类号: Q948. 1



文献标识码 : A



文章编号: 1007-0435( 2009) 01-0131-03
完成时间: 2008 年 6 月
收稿日期: 2008-12-05; 修回日期: 2008-12-08
基金项目:国家
973
项目( 2007CB106801)
作者简介:张红香 ( 1981-) ,女,汉族,辽宁省大连人,博士,研究方向为恢复生态和种子生态学, E- mail: z han ghongxiang8@ hotmail. com
导师简介:周道玮( 1963- ) ,男,汉族,山东青岛人,研究员,主要从事草地饲养及草地管理研究, E-mail: choudaow ei@ yahoo. com. cn
草
地
学
报 第 17卷
Research on Seed Germination Ecology
Candidate: ZHANG Hong-x iang
1
Supervisor: ZHOU Dao-w ei2
( 1. Laboratory of Quant itat ive Vegetation Ecology, Inst itute of Botan y, Chines e Academy of Sciences, Beijin g 100093, China;
2. Northeas t Inst itute of Geography an d Agroecology, Chinese Academ y of S cien ces , Ch angchun, Jilin Province 130012, China)
Life history st rategy is an important resear ch area of plant populat ion ecolog y and seed germ inat ion is
the crit ical stag e of plant life history. T herefore, researches on seed germination r ule and st rateg y help to
know and elucidate the evolut ion and eco logical adaptation t rait s of plant species. P lants in Songnen g rass-
land ar ea w ere t reated as the targ et to invest igate the seed germ inat ion responses to constant temperature
and diurnal increasing o r decreasing temperature, the effect o f global w arm ing simulated by alternat ing
temperature regimes on seed germinat ion of C3 and C4 species, and the germ inat ion pat terns and their dy-
namics under dif ferent temperatures. The germ inat ion response mechanisms of plants to NaCl and PEG
and their interact ion w er e also studied.
Based on the results of constant temperature ef fect, w e pr opo sed the modified thermal t ime model ac-
cording to the basis of thermal t ime model and recruited an equation describing the constant g erminat ion
rate. Germination rate of C3 species w as obviously low er than that of C4 species. Accor ding to thermal t ime
model, the average base germ inat ion temperature o f C3 species w as low er than that of C4 species, but not
significant ly. T he average thermal time constant of C3 species w as higher than that of C4 species and the
dif ference approached to significant level. In this study, plant seed germ inat ion pat tern w as divided into
five categ ories: quick germination, delay ed germination, steady germ inat ion, normal germ inat ion, and de-
layed-quick germinat ion. M ost annual species belonged to the quick germ inat ion pat ter n, w hile the peren-
nial C3 species in meadow belonged to the delayed-quick germinat ion pattern, normal germinat ion pat tern,
or steady germination pat tern. Being an adapt ive st rategy to environments, the seed germinat ion pat tern of
most species changed along w ith different temperatur es. Under global w arming condit ions simulated by a-l
ternat ing temperature reg ime, the germination t rait s o f C3 and C4 species changed. C3 species w as sensit ive
to high temper ature and C4 species sensitive to low temperature. By the germinat ion comparison of all the
alternating temperature reg imes, w e found that sect ional high or low temperature, not the dif ference of
temperature, play ed an impo rtant role in the impacts of g lobal w arm ing on seed germ ination. F ield exper-i
ment results indicate that more plants g erminated in spring in stead of autumn in Songnen grassland. In-
door germination experiment results proved that the germinat ion rate under diurnal incr easing temperature
w as signif icant ly higher than that o f diurnal decreasing temperature w ithin the small temperature range of
spring ( 5~ 10
) and autumn ( 10~ 15
) . Germ inat ion in spring could be an adapt ive st rateg y of plants
from natural select ion.
The relat ionship betw een germ inat ion rate and salinity under dif ferent temperatures w as cur vilinear
fo r Chlor is vi rgata Sw artz. and Dig itar ia sanguinal i s ( L. ) Scop. Some physio logical mechanisms might
be employed by plants to reduce the o smo tic st ress under the m iddle and high salinity conditions leading to
the obviously increase of the germ inat ion r ate. Further experiments show that these tw o species could ger-
m inate and germ inate faster in low er w ater potent ial condition induced by NaCl in comparison w ith PEG
tr eatment . The dry w eight of seeds decreased w ith the t ime pr oceeding . The water content , seed sodium
concentrat ion, and seed solut ion sodium concentrat ion of g erm inated seeds t reated by NaCl w ere signif-i
cant ly higher than those t reated by the iso tonic PEG. Seed sodium concentrat ion and seed solut ion sodium
concentrat ion o f germinated and non-germinated seeds in NaCl t reatments increased along w ith the decrea-
sing of w ater potent ial. Seed w ater content , seed sodium concentrat ion, and seed so lut ion sodium concen-
132
第 1期 张红香:种子发芽生态研究
t rat ion increased linearly along w ith the t ime pr oceeding .
According to the results, w e proposed a salinity model:
S = ( Sm- S) t, in w hich
S is a salinity con-
stant , Sm is the max imum salinity allow ing the seed germination, S is the external salinity, and t is the ger-
m inat ion t ime fo r a given salinity. Seed germination response to salt w as div ided into four stag es. T he f ir st
stage w as mainly involv ed the negat ive o smo tic effect ; at the second stag e, ionic effect and osmot ic effect
ex isted to gether w ith the ionic ef fect st ronger; at the third stage, no seed germinated in PEG treatment
and ionic posit ive effect w as signif icant ly higher than o smo tic negat ive ef fect; at the fourth stag e, the ionic
ef fect began to harm the seed gr adually .
In summary , the results of this study has the theoret ical signif icances and pract ical v alue for the pre-
dict ion of plant germination and emer gence and g lobal w arm ing impacts on plant g row th and dist ribut ion
character ist ics as w ell as the exploitat ion and utilizat ion of w ild plant r esources.
Key words: Seed; Germ inat ion; Temper ature; Thermal time model; Salt ; Osmot ic ef fect; Ionic effect
(责任编辑
李
扬
李
平)
(上接第 127页)


细切捆包青贮与常规捆包青贮一样, 具有作业
效率高、人力负担小、作业方便灵活(不受时间和材
料量限制)、流通方便、避免好气变质等优点。另外,
通过细切,不但提高了饲料的发酵品质,也大大增加
了捆包密度,提高单位容积运输的干物质重量,作为
商品更易流通; 细切捆包更容易解体,解体后不再需
要切短,比常规捆包省时省力; 细切打捆机能捆包
TMR等长度较小的饲料, 用途较广。由于具有上
述优势,细切捆包新技术体系必将得到更大范围的
推广利用,将成为未来青贮的主要方式。
参考文献
[ 1]
M cDonald P, Hender son A R, H erson S . T he biochemist ry of
s ilage [ M ] . M arlow : Chalcomb e Publicat ions, 1991
[ 2]
玉柱,贾玉山,张秀芬.牧草加工贮藏与利用[ M ] .北京:化学工
业出版社, 2004
[ 3]
Wilkin son, J M , T oivonen M I. World silage [ M ] . Lin coln:
C halcombe Publ . , 2003
[ 4]
白春生,玉柱,薛艳林,等.裹包层数对苜蓿拉伸膜裹包青贮品
质的影响[ J] .草地学报, 2007, 15( 1) : 39- 42
[ 5]
崔国文,徐春阳,刘护国,等.紫花苜蓿半干捆包青贮技术的研
究[ J ] .中国草地, 2005, 27( 4) : 15- 19
[ 6]
陶莲,玉柱,陈燕,等. 不同添加剂对红三叶青贮品质的影响
[ J ] .草地学报, 2008, 16( 5) : 522-525, 529
[ 7]
陈娟娟,玉柱,薛艳林,等.添加剂对羊草青贮发酵品质和体外
消化率的影响[ J] .草地学报, 2007, 15( 3) : 238-242
[ 8]
Borreani G, T abacco E. Th e ef fect of a baler chopping system
on fermentat ion and los ses of w rapped big bales of alfal fa [ J ] .
Agronomy Journal, 2006, 98: 1-7
[ 9]
赵毅彬,张勇,何伟宁,等.玉米秸秆打捆青贮机的研究与探索
[ J] . 江西饲料, 2005, 5: 21-24
[ 10]
志藤博克. 細断型
ー
 ー
体系
現状
今後
展開.
自給飼料品質研究会飼料[ J] , 2004, 8-16
[ 11]
野中和久,青木康浩,小林良次, 等.
 ! ∀ # 細断
ー

 ー
 !ー
貯蔵
 品質
 ! ∀ [ J] .日本
草地学会誌 51(別) , 2005, 138-139
[ 12] 塩谷繁,野中和久,西田武弘, 等.細断型
ー
 ー
 調製

長期貯蔵
 ! ∀# ∃ % & ∋ (ー
飼料特性
泌乳
効果[ J] .畜産草地研究成果情報, 2006, 5: 57-58
[ 13] 野中和久,張建国,小林良次, 等.
 ! ∀ # 細断
ー

ー
 !ー
抜
取
調査個数[ J ] . 畜産草地研究成果
情報, 2006, 5: 37-38
[ 14] 河本英憲, 魚住順, 出口新.細断型
ー
 ー
乾物率
25%以上
排汁
低減
重量
推定
可能[ J] . 畜産草地研
究成果情報, 2006, 5: 177-178
[ 15] 喜田環樹,松尾守展,重田一人,等.細断型
ー
 ー


飼料
 高密度成型[ J ] .畜産草地研究成果情報, 2006,
5: 55-56
[ 16] 河本英憲,大谷隆二,山口弘道,等. 細断型
ー
 ー
 活
用
 予乾体系
 ! 稲発酵粗飼料
品質向上技術 [ J ] .
日本草地学会誌, 2005, 51( 2) : 199-201
[ 17] 青木康浩,大下友子,秋山典昭,等.細断型
ー
 ー
 活
用
 高水分
 !∀ # ∃ 排汁抑制技術[ J] . 畜産草地研
究成果情報, 2006, 5: 159-160
(责任编辑
才
杰)
133