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1、4 种群及其基本特征Population and Its Basic Characteristics,4.1 种群的概念 The concept of population4.2 种群动态 Population dynamics4.2.1 种群的密度和分布 Population density and distribution4.2.2 种群统计学 Population demography4.2.3 种群的增长模型 Population growth model4.2.4 自然种群的数量变动 Fluctuation of natural populations,4 种群及其基本特征Popul
2、ation and Its Basic Characteristics,4.2.5 生态入侵 Ecological invasion4.3 种群调节 Population regulation 4.3.1 外源性种群调节理论 Exogenous population regulation theory4.3.2 内源性种群调节理论 Endogenous population regulation theory4.4 集合种群动态 Metapopulation dynamics4.4.1 概念和术语 Concepts and terms4.4.2 集合种群理论的意义与应用 Significanc
3、es and applications of metapopulation theory,教学目的了解种群及其有关名词概念；了解研究种群动态的基本方法；种群密度及其相关概念；种群的内分布型类及相关概念；掌握种群的群体特征及其相关概念；掌握种群增长模型(两种);并学会用两种种群增长模式方程式进行计算；了解自然种群数量变动的表现形式；了解有关种群调节理论的不同学派及其各学派强调的种群调节机制；了解集合种群概念。教学重点与难点1、种群的内分布型;2、种群重要的群体特征;3、种群的增长模型及其应用;4、难点是生命表、出生率、死亡率、生命期望、内禀增长率、生殖价、最小可存活种群、生态入侵的概念.学时数
4、3-4,4.1 种群的概念The concept of population,种群population:同一时期内占据一定空间的同种生物个体的集合。A population is a group of organisms of the same species which occupies a given area.种群边界的不确定性 The boundaries between populations can be arbitrary(任意的).Populations may be categorized as consisting of either unitary or modular
5、organisms.单体生物(unitary organism)和构件生物(modular organism)：In unitary populations,each zygote受精卵 gives rise to a single individual.In modular organisms,the zygote develops into a unit of construction which gives rise to further modules and a branching structure.The structure may then fragment producing
6、 many individual ramets无性系分株.,构件生物,由一个合子发育的形成一套构件组成的个体。如一株树有许多树枝。并且构件数很不相同，随着环境的变化而变化。一般高等植物是构件生物，大多数动物属单体生物。但营固着生活的如珊瑚，薮枝虫、苔藓也是构件生物。One that grows by the repeated iteration(迭代)of parts,e.g.the leaves,shoots and branches of a plant,the polyps(个员)of a 珊瑚coral or 苔藓虫bryozoan.Modular organisms are almo
7、st always branched,though the connections between branches may separate or decay and the separated parts may in many cases then become physiologically independent,e.g.水螅Hydra spp.and 浮萍duckweeds(Lemna spp.).,构件生物,苔藓虫bryozoan：,无性系分株ramet,构件生物各部分间的连接可能会死亡和腐烂，这样就形成了许多分离的个体，这些个体来源于一个受精卵并且基因型相同。,种群的3个基本特
8、征Three basic characteristics of a population,空间特征Spatial characteristics：一定的分布区域数量特征Numerical characteristics：变动的个体数量遗传特征Genetic characteristics：一定的基因组成,4.2 种群动态 Population Dynamics,种群数量的时空变动规律Spatio-temporal variation law of population size1)how many:数量number和密度density2)where:分布distribution3)how:数量
9、变动fluctuation和扩散迁移diffusion4)why:种群调节regulation基本研究方法：野外调查field investigation、实验验证experimental validation、模型模拟modeling and simulation,4.2.1 种群的大小和密度,大小size:一定区域的个体数量、生物量或能量密度density:单位面积（单位体积、单位生境）中的个体数目区分单体生物和构件生物：如水稻分蘖。绝对密度和相对密度种群密度的估计方法：总数量调查法、样方法、标记重捕法（N:M=n:m),Population Size,The population siz
10、e for unitary organisms,such as mammals,is simply the number of individuals in a give area.For modular organisms,such as plants and corals,the situation is more complex.In this case the number of pieces(ramets)or the number of shoots(modules)may give a more meaningful indication of abundance than th
11、e number of different individuals.,4.2.1.3 种群的空间结构Spatial Structure,分布格局distribution pattern:The spatial distribution at any particular moment of the individuals of a species of plant or animal.组成种群的个体在其生活空间中的位置状态或布局。参见：戴小华,余世孝.GIS 支持下的种群分布格局分析J.中山大学学报(自然科学),2003,42(1):75-78,分布格局,3种：随机random、成群clump
12、ed、均匀uniform,Antagonistic 敌对的，depletion 枯竭,演替进程中的分布格局变化,4.2.2 种群统计学 Demography,种群基本特征：种群密度初级种群参数：出生率natality、死亡率mortality、迁入immigration、迁出emigration；与种群密度变化密切相关；最大出生率、实际出生率、特定年龄出生率；最低死亡率、生态死亡率、特定年龄死亡率次级种群参数：性比、年龄结构、种群增长率,Concepts,Natality is the birth of new individuals.The realized natality实际出生率 is
13、 the actual successful reproduction per female over a period of time.The age-specific birthrate特定年龄出生率 is the number of offspring produced per unit time by females in specific age classes.Mortality:The death rate,or mortality rate,is the number of individuals dying during a given time interval divid
14、ed by the average population size over that time interval.This is an instantaneous(瞬时的)rate and can be estimated for the population as a whole or for specific age classes to give the age specific mortality rate特定年龄死亡率.The probability of dying is the number dying per individual present at the start o
15、f the time period,4.2.2 种群统计学 Demography,4.2.2.1 年龄、时期结构和性比 Age,stage structure and sex ratio4.2.2.2 生命表、存活曲线和种群增长率 Life table,survivorship curve and population growth rate 1 生命表 2 K-因子分析 K-Factor analysis 3 存活曲线 4 种群增长率和内禀增长率intrinsic natural rate of increase 5 生殖价 reproductive value,4.2.2.1 年龄、时期结
16、构和性比,年龄椎体：age pyramida 增长型种群b 稳定型种群c 下降型种群,4.2.2.1 年龄、时期结构和性比,年龄结构：一个年龄群对整个种群的比率时期结构stage structure：适用于离散发育期的生物如昆虫个体大小群size classes：适用于生长率与年龄无密切关系的生物如植物性比sex ratio：种群中雌雄个体的比例。多数=1：1；孤雌1；社会昆虫1；性比可随环境变化；性转变,Age and stage structure,The age structure of a population is the number of individuals in each
17、age class expressed as a ratio,and is usually displayed as an age pyramid diagram.A population which is neither expanding nor contracting will have a stationary age distribution.A growing population will have more young,while a declining population will be dominated by older age classes.Where organi
18、sms pass through discrete growth stages(e.g.insect larval instars幼虫龄期),the number of individuals at each stage(the stage structure)may provide a useful description of the population.In species where growth rates are indeterminate(不确定,such as plants),size classes may be more informative.,性转变,性别转变现象可以
19、说是行为生态学中最有趣、最奇异的现象之一。在雄性动物之间只要存在激烈的择偶竞争，通常是只有个体最大和最强壮的雄性才能占有最大的生殖优势，而小者或弱者为了回避和强大对手的直接竞争往往采取偷袭交配的对策。但是，一个更令人吃惊的对策就是改变性别，即借助于性别的转化改变自己的不利处境，以获得生殖上的较大成功。,性转变,雌性变雄性往往是当动物还没有充分长大时，它先作为一个雌性个体参与繁殖；但当它一旦长大到足以赢得竞争优势的时候便转变为雄性，开始以雄性个体参与繁殖。这种情况在鱼类中是比较常见的。性别发生转变往往比终生保持一种性别能在生殖上获得更大的好处，因为对改变性别的个体来说，它无论是在小而弱时，还是在
20、大而强时，都能得到生殖的机会。就其一生的生殖来说，改变性别的个体也比不改变性别的个体更为成功。这个问题与种群的性比率问题是密切相关的，它们都是资源在两性间如何分配这一大问题中的一个局部问题。较为少见的现象是，当动物个体较小时表现为雄性，一旦长大后便转变为雌性。这种类型的性别转变只有当雄性动物之间的择偶竞争不太激烈和雄体大小对生殖成功又无太大影响时才有利。由于择偶竞争不激烈，所以小个体的雄性动物也能使一些生殖力较强的大个体雌性动物受精。,生命表 Life Table,种群统计的核心是建立反映种群全生活史的各年龄组或生活态级出生率、死亡率，甚至包括迁移率在内的信息综合表。,一般的人口生命表格式或构
21、成，表头依序是：,x：年龄级nx:在x龄级开始时的存活个体数lx:在x龄级开始时的标准化的存活个体数dx:从x到x+1期标准化的死亡个体数qx:从x到x+1期的死亡数ex:x期开始时的平均期望寿命或平均余年Lx:从x到x+1龄期的平均存活个体数Tx:龄期x及其以上各年龄级的个体存活总年数,Life Tables,Life tables summarize the fate of a group of individuals born at approximately the same time from birth to the end of the life cycle.Such a gro
22、up is known as a cohort同生群 and investigation of this kind is termed cohort analysis.Life tables show the number of individuals present at different life stages or ages together with age-specific survival rates and age-specific mortality rates calculated from each stage.Mortality at each stage is exp
23、ressed by k-values which are derived from logarithms and can be summed to give total mortality.,藤壶的生命表,Lx：在 x 期到x+1 期间平均存活数目=(lx+lx+1)/2 Tx：在 x 期限后的平均存活数的累计数=Lx,K-因子分析 K-Factor Analysis,A statistical treatment of population data designed to identify factors most responsible for change in population
24、size.The technique allows that the identification of key factors contributing to mortality.Stage-specific k-values obtained over successive years are compared to the values for total mortality(ktotal).K-Factor analysis highlights those stages suffering the greatest mortality which are responsible fo
25、r fluctuations in loss rate and hence population size.,microsporidian 小孢子虫的,存活曲线 Survivorship Curve,Survivorship is the converse of mortality.Survivorship data are often shown as a survivorship curve for a particular population;a graph showing the proportion of survivors on a logarithmic scale throu
26、gh each phase of life.There are three generalized patterns of age-specific survivorship depending on whether the probability of dying is highest later in life(Type I),constant through life(Type II)or highest for young stages(Type III).,存活曲线 Survivorship Curve,I型：凸型，大型哺乳动物、人类；II型：对角线型，一些鸟类；III型：凹型，产卵
27、鱼类、贝类和松树,Dall sheep野大白羊、戴尔绵羊,种群增长 Population Growth,r 种群的实际增长率即自然增长率rm 种群的内禀增长率 intrinsic natural rate of increase:This is the theoretical maximum that may be reached in a given environment if the population is not resource-limited.b 出生率d 死亡率,种群增长 Population Growth,R0 世代的净增殖率 basic reproductive rate
28、:This is the number of offspring produced per original individual by the end of the cohort.In an annual population,it indicates the overall extent to which the population has increased or decreased over that timeT 世代时间！计划生育的生态学基础计划生育的种群生态学基础：要降低r，要么降低R0（少生），要么增大T（晚婚晚育）,tunicate 被囊类动物,内禀增长率 innate ra
29、te of increase,内禀增长率rm：按Andrewartha和Brich的定义（1954），内禀增长率是指具有稳定年龄结构的种群，在食物与空间不受限制、同种其他个体的密度维持在最适水平、环境中没有天敌、并在某一特定的温度、湿度、光照和食物性质的环境条件组配下，种群的最大瞬时增长率。反应了种群在理想状态下，生物种群的扩繁能力。,生殖价 Reproductive Value,生殖价Vx：用来描述某一年龄的雌体评价能对未来种群增长所做的贡献。是衡量种群内个体繁殖力和存活力的一个综合指标，对生活史性状的进化非常重要：,4.2.3 种群的增长模型Population Growth Model,
30、4.2.3.1 与密度无关的种群增长模型 Density-independent population growth models 1 种群离散增长模型 Discrete models of population growth 2 种群连续增长模型 Continuous models of population growth4.2.3.2 与密度有关的种群增长模型 Density-dependent population growth models 逻辑斯谛方程 Logistic equation,与密度无关的种群增长模型,种群离散增长模型几何增长Geometrical growth种群连续增
31、长模型指数增长 Exponential growth连续增长种群加倍时间,phlox 福禄考属,种群连续增长模型,Scots pine欧洲赤松,collared dove斑鸠,全球人口增长曲线,逻辑斯谛方程 Logistic Equation,逻辑斯谛方程 Logistic Equation,rinder pest牛瘟,牛疫,淡水枝角水蚤(Daphnia pulex),逻辑斯谛方程的重要意义,许多两个相互作用种群增长模型的基础;渔业、牧业、林业等领域确定最大持续产量的主要模型;模型中两个参数r 和K，已成为生物进化对策理论中的重要概念。,4.2.4 自然种群的数量变动,4.2.4.1 种群增长
32、：J型、S型和过渡中间型4.2.4.2 季节消长Seasonal change：4.2.4.3 种群的波动Population fluctuation：环境随机变化、时滞密度制约、过偿性密度制约 1 不规则波动；2 周期性波动。4.2.4.4 种群的爆发Population explosion：4.2.4.5 种群平衡Population equilibrium：4.2.4.6 种群的衰落与灭亡Population decline and extinction：4.2.4.7 生态入侵Ecological invasion：,Fluctuation,The abundant rains of
33、1983(a)greatly increased plant growth on the Galpagos Islands compared to(b)periods of lower rainfall,Rainfall and the medium ground finch,Geospiza fortis,population of Daphne Major Island(data from Gibbs and Grant 1987).,生态入侵的实例,原产于澳大利亚的桉树是经常被大量引种的外来物种之一，却屡屡引起诸如土壤贫瘠、地下水位下降和生物多样性降低等严重的生态问题水葫芦（Eichho
34、rnia crassipes）主要是作为饲料被引入到我国，也作为观赏和净化水质的植物得到推广种植。过去10年，广东、云南、江苏、浙江、福建、上海等省市每年都要人工打捞水葫芦，仅浙江温州市和福建莆田市1999年用于人工打捞水葫芦的费用分别为1000万元和500万元，全国总的费用有多少，目前没有准确统计，至少超过1亿元，而水葫芦带来的农业灌溉、粮食运输、水产养殖以及旅游等方面的经济损失更大。福寿螺、罗非鱼、紫茎泽兰、大米草、薇甘菊等,4.3 种群调节,4.3.1 外源性种群调节理论：强调外因4.3.1.1 非密度制约的气候学派：强调气候，研究对象多为昆虫4.3.1.2 密度制约的生物学派：强调捕食
35、、寄生和竞争等生物过程折中观点4.3.2 内源性自动调节理论：强调内因4.3.2.1 行为调节温-爱德华学说：社群行为4.3.2.2 内分泌调节克里斯琴学说：主要适用于兽类4.3.2.3 遗传调节奇蒂学说,4.4 集合种群动态,集合种群metapopulation：A population perceived to exist as a series of subpopulations,linked by migration between them.However,the rate of migration is limited,such that the dynamics of the m
36、etapopulation should be seen as the sum of the dynamics of the individual subpopulations.生境斑块中局域种群(local population)的集合，这些局域种群在空间上存在隔离，彼此间通过个体扩散而相互联系，也有人成为一个种群的种群(a population of populations),集合种群类型,4.4.2 集合种群理论的意义与应用,景观管理和自然保护1）大范围害虫防治问题；2）物种灭绝概率与斑块面积的关系；3）人工迁移或走廊防止物种灭绝4）SLOSS争论：一个大保护区还是多个相互联系的小保护区
37、？The SLOSS Debate was a debate in ecology and conservation biology during the 1970s and 1980s as to whether a Single Large or Several Small(SLOSS)reserves was a superior means of conserving biodiversity in a fragmented habitat.,参考书目,1 张大勇.2000.理论生态学研究.北京：高等教育出版社，施普林格出版社2 蒋志刚等.2000.保护生物学.杭州：浙江科学技术出版社3 孙儒泳.2001.动物生态学原理.北京：北京大学出版社4 Mackenzie A et al.2003.Instant Notes in Ecology.北京：科学出版社版权申明：本课程图片除本人拍摄外，均来自互联网和Molles J M C 2002，仅用于教学目的（来源已在图片格式的web可选文字中注明）,作业,p852.试说明我国计划生育政策的种群生态学基础。3.Logistic equation的模型式、各参数意义、以及该方程的生态学涵义。,
