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Essentials of Ecology
鄭先祐 (Ayo)
國立臺南大學 環境與生態學院
生物科技學系 生態學 (2008)
教科書
• Essentials of Ecology,
3rd Edition
• Townsend, Begon, Harper
• March 2008, Wiley-Blackwell
• ISBN: 978-1-4051-5658-5
Website: http://www.blackwellpublishing.com/townsend/
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About the Authors
• Colin R. Townsend is the Director of the Ecology,
Conservation and Biodiversity Research Group at
the University of Otago. He has published over 150
ecology papers and books, including the bestselling and award-winning Ecology, 4th Edition, with
co-authors Michael Begon and John L. Harper. He
is also co-editor of the international journal
Freshwater Biology.
• Michael Begon (University of Liverpool) (animal
population ecology)
• John L. Harper (University of Exeter) (plant population
biology)
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• Colin R. Townsend (University of Otago)
(conservation biologist)
• Michael Begon is Professor of Ecology in the
School of Biological Sciences at the University of
Liverpool. He has authored several books in
addition to Ecology and around 150 research
articles. His current research interests are focused
on the population ecology of pathogens in wildlife
populations.
• John L. Harper is an award-winning biologist and
a leading figure in plant population biology. He has
served as a council member of the Fellowship of
the Royal Society and received the Darwin Arward
in 1990. He is also the author of Population
Biology of Plants and a co-author of Ecology.
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Contents
•
Preface
•
Acknowledgments
• Part I: Introduction: (簡介)
• Part II: Conditions and Resources: (情境與資源)
• Part III: Individuals, Populations, Communities and
Ecosystems: (個體、族群、群落與生態體系)
• Part IV: Applied Issues in Ecology: (應用生態學)
•
References
•
Index
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Contents
• Part I: Introduction:
– 1. Ecology and how to do it (簡介)
– 2. Ecology‘s evolutionary backdrop (演化背景)
• Part II: Conditions and Resources:
– 3. Physical conditions and the availability of
resources (物理情境與資源的供應力)
– 4. Conditions, resources and the world‘s
communities (情境、資源和世界群落)
• Part III: Individuals, Populations,
Communities and Ecosystems:
• Part IV: Applied Issues in Ecology:
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Contents
• Part I: Introduction:
• Part II: Conditions and Resources:
• Part III: Individuals, Populations, Communities
and Ecosystems:
–
–
–
–
–
–
–
5. Birth, death and movement (出生、死王和遷移)
6. Interspecific competition (種間競爭)
7. Predation, grazing and disease(掠食、草食和疾病)
8. Evolutionary ecology (演化生態學)
9. From populations to communities (族群到群落)
10. Patterns in species richness (物種豐富度)
11. The flux of energy and matter through ecosystems
(生態体系的能量與物質流動)
• Part IV: Applied Issues in Ecology:
7
Contents
•
•
•
•
•
Preface
Acknowledgments
Part I: Introduction:
Part II: Conditions and Resources:
Part III: Individuals, Populations, Communities
and Ecosystems:
• Part IV: Applied Issues in Ecology:
– 12. Sustainability (可持續力)
– 13. Habitat degradation (棲地衰退)
– 14. Conservation (保育)
• References
• Index
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Ecology: concepts & Applications
Molles, M. C. Jr. (2008) 4th ed. McGraw-Hill Co., Inc.
•
•
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Introduction: What Is Ecology? (chap.1)
Section I Natural History (chap. 2, 3)
Section II Individuals (chap. 4,5,6,7)
Section III Population Ecology (chap.
8.9.10.11.12)
• Section IV Interactions (chap.13,14,15)
• Section V Communities and Ecosystems
(chap.16,17,18,19,20)
• Section VI Large-Scale Ecology (chap. 21,22,23)
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Fundamentals of Ecology
By Odum, E. P. and G. W. Barrett (2005)
1. The Scope of Ecology
2. The Ecosystem
3. Energy in Ecological systems
4. Biogeochemical Cycles
5. Limiting and Regulatory Factors
6. Population Ecology
7. Community Ecology
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Fundamentals of Ecology
8. Ecosystem Development
9. Landscape Ecology
10. Regional Ecology: Major Ecosystem Types and
Biomes
11. Global Ecology
12. Statistical Thinking for Students of Ecology
•
Glossary
•
References
•
Credits
•
index
11
Odum, E. P. (1997) Ecology: A bridge
between Science and Society. Sinauer
Associates, Inc.
Brief Contents
•
•
•
•
•
•
•
•
•
•
緒言：Apollo 13號的航行
第一章 維生環境
第二章 組織的層次
第三章 生態體系
第四章 能量學
第五章 生存的物質循環和物理環境
第六章 族群與群區生態學
第七章 發展與演化
第八章 世界主要生態體系的類型
結語：從年輕至成熟的轉變
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Preface
• 這本書的目的：
– 分享自然複雜世界的美妙
– 面對受傷害的自然，我們大家都需要有生態學
識，才有能力為保護自然辯護，以及貢獻於化
解生態問題。
• 這本書的適用：
– A semester-long beginners course
• 完整的生態學：可參閱
– Ecology: from individuals to ecosystems
(Begon, Townsend & Harper, 4th ed. 2005)
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參考書
• Ecology: From Individuals to
Ecosystems, 4th Edition
• Begon, Townsend, Harper
•
•
•
•
ISBN: 978-1-4051-1117-1
Paperback
752 pages
August 2005, Wiley-Blackwell
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Essentials of Ecology 3rd. Ed.
Chap. 01 Ecology and how to do it
鄭先祐 (Ayo)
國立臺南大學 環境與生態學院
生物科技學系 生態學 (2008)
Contents
Part I: Introduction:
• Chap.1. Ecology and how to do it
– 1.1 Introduction
– 1.2 Scales, diversity and rigor (尺度、多樣性、精確)
– 1.3 Ecology in practice (生態學的研究)
• Chap.2. Ecology's evolutionary backdrop
Part II: Conditions and Resources:
Part III: Individuals, Populations,
Communities and Ecosystems:
Part IV: Applied Issues in Ecology:
16
Key concepts
1. Define ecology (an applied and a pure science)
2. To describe and understand, and on the basis
of their understanding, to predict, manage, and
control
3. Spatial and temporal scales(時空尺度). Patterns
may be evident only at particular scales
4. By means of observations, field and laboratory
experiments, and mathematical models.
5. Relies on truly scientific evidence (and the
application of statistics)
17
1.1 introduction
• Ecology (originally in German, Oekologie),
was first defined in 1866 by Ernst Haeckel.
– Comprehensive science of the relationship of
the organism to the environment.
– The science which concerns itself with the
external relations of plants and animals to
each other and to the past and present
conditions of their existence, to be contrasted
with physiology (internal relations) and
morphology (structure). (Burdon-Sanderson,
1893)
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「生態學」的定位(起源)
1866-1891 Haeckel, Ernst:
第一個使用 ecology (1866)。
– Ecology as a part of physiology (生理學)。
– total relationships of the animal both to its
inorganic and its organic environment.
– complex interrelations referred to by Darwin as
the conditions for the struggle for existence.
• eco-logy (the study of household)
– 字源取自 eco-nomics (household + management)
– the economy of nature (自然的經濟)
19
「生態學」的定位( by 1920s)
• 1893 J. S. Burdon-Sanderson: President,
British Ass. for Advancement of Science.
– 將 Ecology 成為「生物學」中三大部之一。
– ecology, physiology and morphology
– 生態學 => philosophy of living nature.
• 1900s Clements & Cowles → plant ecology
– 生態學 = 生理學
• 1910s Adam (1917) & Elton (1927) → animal ecologists
– 生態學= new natural history (Adam, 1917)
= scientific natural history (Elton, 1927) ┤
20
「生態學」的定位( by 1950s)
• 1949 Allee et al. principles of animal ecology
– lack of knowledge of ecology among scholars and
philosophers
• 1. 由於內在的多元性 → 廣泛 distortion of 內容
• 2. 缺乏 historical studies
21
「生態學」的定位( 1960-80s)
• 生態學 =
– 生物的分佈和豐富量之研究 (Andrewartha, 1961)
– 自然的結構和功能之研究 (Odum, 1963)
– 「生物與其生活環境」的互動關係之研究 (Leo
Smith, 1986)
– 自然科學和社會科學的橋樑 (Odum, 1983)
22
「生態學」的定位( 1990s)
• 生態學 =
– 五部份：演化生態+行為生態+族群生態+群落
生態+應用生態 (Stiling, 1992)
– 主要兩個根源：植物地理學 和 自然歷史學
(Leo Smith, 1992)
– the Science of the universe(宇宙) (Hutchinson)
(Colinvaux, 1993)
•
•
•
•
1. 体系的龐雜和歧異之維持和來源之瞭解
2. 生物一起生活的原則
3. 生命如何受天擇影響
4. 自然的經濟學
23
「生態學」的定位( by 2000)
• 生態學 =
– 生命與維生環境之關係 (Odum, 1993)
– 決定「生物的分佈和豐富量」之互動的研究：
Where, How many, Why (Krebs, 1994)
– 「生物與其生活環境」的互動關係之研究
(Brewer, R., 1994)
– 生物與生活環境間的互動關係。 (Stiling, P.
D.,1996
– 自然的經濟 (Ricklefs, R. E.,1997; Smith and
Smith, 1998)
– 研習自然世界的次序 (order) (Ricklefs and
Miller,2000)
24
Charles Elton (1900-1991), Animal Ecology (1927)→
Essentials of Ecology (3rd. Ed.)
• The scientific study of the distribution and
abundance of organisms and the interactions
that determine distribution and abundance.
(Townsend, Begon and Happer, 2008)
• The primitive humans must have been
ecologists of sorts – driven by the need to
understand where and when their food and
their enemies were to be found. (applied
ecologists).
25
Fundamental problems
• To what extent does competitions for food
determine which species can coexist in a habitat?
• What role does disease play in the dynamics of
populations?
• Why are there more species in the tropics than at
the poles?
• What is the relationship between soil productivity
and plant community structure?
• Why are some species more vulnerable to
extinction than others?
26
Essentials of Ecology (3rd. Ed.)
• The beauty of ecology is that it challenges
us to develop an understanding of very
basic and apparent problems – in a way
that recognizes the uniqueness and
complexity of all aspects of nature – but
seeks patterns and predictions within this
complexity rather than being swamped by it.
27
Essentials of Ecology (3rd. Ed.)
•
Two different classes of explanation in biology
– Proximate explanation
– Ultimate explanation
•
學習研究的階段
– 描述、機制(功能)、模式、預測 (參閱實習手冊)
•
This book is about:
1. How ecological understand is achieved.
2. What we do understand.
3. How that understanding can help us predict, manage
and control.
28
「生態學」的定位
生態學 = Eco + logy = 生活科學 =
自然的經濟 (Economy of Nature)
經濟學(Economics) =Eco + nomics
= 生活的經營管理
• 生態學 (Ecology)
– 生活科學：研究生物生活的學
理 (生存與發展之道)
• 生物：個體、族群、群落，以及
生態體系
• 生活：存活 + 繁殖 (生生不息)
29
經濟學 vs.生態學
Odum, E. P. and G. W. Barrett (2005) Fundamentals of Ecology.
30
1.2 Scales, diversity and rigor
(尺度、多樣性、精確)
Three points
1. Ecological phenomena occur at a variety
of scales
2. Ecological evidence comes from a variety
of different sources
3. Ecology relies on truly scientific evidence
and the application of statistics.
31
Chap. 1 Ecology and how to do it
• 1.1 introduction
• 1.2 Scales, diversity and rigor
– 1.2.1 Questions of scale
– 1.2.2 the diversity of ecological evidence
– 1.2.3 statistics and scientific rigor
• 1.3 Ecology in practice
– 1.3.1 Brown trout in New Zealand
– 1.3.2 Successions on old fields
– 1.3.3 Hubbard Brook: a long-term
– 1.3.4 a modeling study: vultures extinction
32
組成的位階
分類的位階
Biosphere
Ecosystem
Clade
Community
複雜
Species
Population
Deme
Organism
Organism
Organ System
1.2.1 Questions of scale
Organ
Tissue
圖a-1. 生物的組成與
分類的位階。clade 包
含：種以上的各個分
類位階。原資料摘取
自Kohn (1989), p.1097。
Cell
Organelle
Macromolecule-gene
Molecule
簡單
33
8
6
地球：生物界
生物學
台灣生態
4
空
1km
2
間
的
生物体
1m
0
小型生物
量
度
大型生物
-2
無脊椎動物
1mm
原生物
-4
胞器
1um -6
細胞
病毒
蛋白質
-8
小分子
1A -10
-16 -14 -12 -10
1 微秒
-8
-6
1分
-4 -2 0 2
1天
1年
4
6
8
10
1 百萬年
時 間 的 量 度
圖a-2. 生物學的時間與空間的範圍。中間方形區是指「生物個体」
的生物學。時間量度以年為單位；空間則以公尺為單位。基本資料
34
取自Kohn (1989), p.1096。
1.2.2 the diversity of ecological evidence
1. Observations and field experiments
2. Laboratory experiments
– Simple laboratory systems
– Mathematical models
35
1.2.3 statistics and scientific rigor
1. Ecology: a search for conclusions in
which we can be confident
2. Ecologists must think ahead
3. Ecology relies on representative samples
36
Species
分
陸域
類
位
海域
階
Phylum
0%
100%
百分率 (%)
圖a-3. 於各個分類位階，陸域與海域生態体系的龐雜
度之比較。資料取自Ray (1985) Amer. Zool., 24:452468。
37
圖a-4. 一般生命的延續
過程。「生命」起自
「基因型」(genotype,
Phenotype space
受精卵, G1)，經過t1時
t2
間，發展成「個体」(表
P1
現型，phenotype，P1)；
再經過t2時間發展成為
t1
「成熟的個體」(P2)，
經過t3時間產生「配子」
(精子或卵子，G2)；再 Genotype space
經過t4時間結合成「受
精卵」(G‘1)，另一個新
生命的開始，進入另一
G1
個循環。
P‘2
P2
t‘2
P‘1
t3
t‘1
t’3
t“1
G‘2
t‘4
G2
t4
G‘1
G“1
38
整合的生物學
--生態体系生物學
--群落(社區)生物學
真菌界
原核類
植物界
病毒類
動物界
原生物
族群生物學
個体生物學
生理與解剖學
組織生物學
細胞生物學
分子生物學
系統分類學
圖a-5. 生物學的結構。
親緣遺傳學
生
物
學
的
根
基
演化生物學
39
• Fig. 1-4
Compared with
the strong setpoint controls at
the organism level
and below,
organization and
function at the
population level
and above.
40
Fig. 1-2. Ecological levels-of-organization spectrum emphasizing the
interaction of living and nonliving components.
41
1.3 Ecology in practice
• 1.3.1 Brown trout in New Zealand: effects
on individuals, populations, communities
and ecosystems
• 1.3.2 Successions on old fields in
Minnesota: a study in time and space
• 1.3.3 Hubbard Brook: a long-term
commitment of large-scale significance
42
1.3.1 Brown trout in New Zealand
• (a) a brown trout (外來種)：視覺覓食
• (b) a Galaxias fish (本土種)：動態感應覓食
• 食物：Mayfly nymphs (浮游蛹)
43
白天量少
白天量少
• Fig. 1.7 (a) Mean
number (+SE) of
mayfly nymphs
collected either from
a trout stream or a
Galaxias stream. (b)
mean number (+SE)
of mayfly nymphs
observed on the
upper surfaces of
cobbles during late
afternoon .
44
45
藻類量多
無脊椎動
物量少
• Fig. 1.8 (a) total invertebrate biomass
• (b) algal biomass (chlorophyll a)
• T, trout present; G, Galaxias present; N, no fish.
46
• Fig. 1.9
47
1.3.1 Brown trout in New Zealand
• Individual level: trout → Mayfly 白天活動量
少
• Population level: trout 影響 Galaxias fish的
族群分布
• Communities (ecosystem) level:
• Brown trout ← invertebrates ← algae
– Energy flow
– Cause a cascade of effects
48
1.3.2 Successions on old fields: a study in
time and space
• Ecological succession
• (a) invader species
decreased
• (b) native prairie
species increased
• (c) annual species
decreased
• (d) perennial species
increased
• (e) soil nitrogen
increased
49
Nitrogen effects
• Do patches receiving different supply rates
of nitrogen become less similar in species
composition over time?
• Do patches receiving similar supply rates
of nitrogen become more similar in
species composition over time?
50
• Fig. 1.11 (a) between 1982 and 1992, plots receiving
17g of nitrogen/m2/yr in the two fields became
increasingly similar in composition.
• (b) with only 1 g of nitrogen/m2/yr.
51
1.3.3 Hubbard Brook: a long-term
commitment
• Fig. 1.12 the
Hubbard Brook
experimental
forest.
• The value of
large-scale
studies and longterm data records
52
• The output of chemicals was greater than
their input from rain, sleet and snow,
except of nitrogen.
53
• Fig. 1.13
deforested 和
control 比較
54
• Fig. 1.14 Longterm changes in
concentrations
• Clean Air Act in
1970, emissions of
SO2 were reduced.
• 但是Ca+也下降。
• Acid rain may be
responsible
• Forest growth
decline.
55
1.3.4 A modeling study: vultures
extinction
• In 1997, vultures in
India and Pakistan
began dropping
from their perches.
• Population surveys
from 2000 to 2003
confirmed alarming
rates of decline.
• =0.50-0.52
Oriental white-backed vulture Gyps bengalensis
56
• Vultures 族群下降，野外腐肉增多，蒼蠅、野狗、
鼠類等增多，傳染病增多。
• 信仰，鳥葬 (a special tower, dakhma)
• It took a few years to find, vultures 死因是累積尿
酸在體內，最後腎臟衰竭。
• 這是因為吃食到 diclofenac 藥物。
• 抗發炎的非類固醇藥物，1970年代使用於人體，後
來於Pakistan和印度廣泛使用於牲畜。
• 但這是唯一的因素嗎？
57
Fig. 1.14
58
59
6 about models
• A model is a formulation that mimics a real-world
phenomenon.
• Computer-simulated models permit one to
predict probable outcomes as parameters in the
model are changed.
• Symbols:
–
–
–
–
E= energy source (forcing function)
P= state variables
F= flow pathways
I=interaction functions
60
F= flow pathways
L= feedback loop
Fig. 1-6 Compartment diagram showing the five basic
components of primary interest in modeling ecological
systems.
61
Fig. 1-6 Compartment model with a
feedback or control loop that transforms a
linear system into a partially cyclical one.
62
隔離
Fig. 1-7 Interaction of positive and negative feedbacks
in the relationships of atmospheric CO2, climate
warming, soil respiration, and carbon sequestration.
63
Fig 1-8. The H. T. Odum energy language symbols used in
model diagrams.
64
Fig. 1-9 Ecosystem model using energy language symbols and
including estimated rates of energy flow for a Florida pine forest.
65
Boxes
• 1.1 historical landmarks
• 1.2 quantitative aspects: interpreting
probabilities
• 1.3 quantitative aspects: attaching
confidence to results
• 1.4 quantitative aspects: estimation:
sampling, accuracy and precision
• 1.5 Topical ECOncerns: invasions and
homogenization of the biota: does it matter?
66
Questions
1. Search the library for a variety of
definitions of ecology: which do you think
is most appropriate and why?
2. Discuss the pros and cons of descriptive
studies as opposed to laboratory studies
of the same ecological phenomenon.
67
問題與討論
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