Ecology 鄭先祐 (Ayo) 教授

國立台南大學 環境與生態學院生態科學與技術學系

環境生態研究所 + 生態旅遊研究所

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生態學 (Ecology) 課程大綱 ( 整體 )

T01. 簡介：生態學 (Chap.1)

I. 個體與環境 (Chap.2,3,4,5,6)

II. 族群生態學 (Chap.7,8,9,10)

III. 個體間互動 (Chap.11,12,13,14)

IV. 群落生態學 (Chap.15,16,17,18)

V. 生態體系生態學 (Chap.19,20,21)

VI. 應用生態學 (Chap.22,23,24)

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生態學 (Ecology) 課程大綱 (上 )

T01. 簡介：生態學 (Chap.1)I. 個體與環境

T02. 物理環境與生物界 (Chap. 2,3) T03. 環境適應 (Chap. 4,5) T04. 演化學與生態學 (Chap. 6)

II. 族群生態學 T05. 生活史與族群分布 (Chap. 7, 8) T06. 族群成長與變動 (Chap. 9,10)

III. 個體間互動 T07. 競爭、掠食與草食 (Chap.11,12) T08. 寄生與互利共生 (Chap.13,14)

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生態學 (Ecology) 課程大綱 (下 )

Unit IV. 群落生態學 T09. 群落與演進 (替 ) (Chap. 15,16) T10. 生物地理 (Chap.17) T11. 物種多樣性 (Chap.18)

Unit V. 生態體系生態學 T12. 生產、能量循流與食物網 (Chap.19,20) T13. 營養供應與循環 (Chap.21)

Unit VI. 應用生態學 T14. 保育生物學 (Chap.22) T15. 景觀生態學與生態體系經營管理 (Chap.23) T16. 全球生態學 (Chap.24)

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T01. 簡介：生態學• Chap.1 Introduction: The web of Life

I. 個體與環境 T02. 物理環境與生物界

• Chap.2 the Physical Environment• Chap.3 the Biosphere

T03. 環境適應 • Chap.4 Coping with Environmental variation:

Temperature and Water• Chap.5 Coping with Environmental variation: Energy

T04. 演化學與生態學• Chap.6 Evolution and Ecology

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II. 族群生態學 T05. 生活史與族群分布

• Chap.7 Life History Analyses• Chap.8 Population distribution and abundance

T06. 族群成長與變動 • Chap.9 Population growth and regulation• Chap.10 population dynamics

III. 個體間互動 T07. 競爭、掠食與草食

• Chap.11 Competition• Chap.12 Predation and Herbivory

T08. 寄生與互利共生 • Chap.13 Parasitism• Chap.14 Mutualism and Commensalism

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Unit IV. 群落生態學 T09. 群落與演進 (替 )

• Chap.15 The Nature of Communities• Chap.16 Change in Communities

T10. 生物地理• Chap.17 Biogeography

T11. 物種多樣性 • Chap.18 Species diversity in communities

Unit V. 生態體系生態學 T12. 生產、能量循流與食物網

• Chap.19 Production• Chap.20 Energy flow and food webs

T13. 營養供應與循環 • Chap.21 Nutrient supply and cycling

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Unit VI. 應用生態學 T14. 保育生物學

• Chap.22 Conservation biology T15. 景觀生態學與生態體系經營管理

• Chap.23 Landscape Ecology and Ecosystem management

T16. 全球生態學 • Chap.24 Global Ecology

Appendix Answers to review questions Glossary Illustration credits Literature cited index

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作者 I ： Michael L. Cain

Ph.D. in Ecology and Evolutionary Biology from Cornell University

Taught at New Mexico State University and the Rose-Hulman Institute of Technology, and is currently affiliated with Bowdoin college.

His research interests include: plant population ecology, long-distance dispersal, ecological and evolutionary dynamics in hybrid zones, and search behavior in plants and animals.

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作者 II ： William D. Bowman

His Ph.D. from Duke UniversityIs a professor at University of Colorado

at Boulder, affiliated with both the Department of Ecology and Evolutionary Biology and the Institute of Arctic and Alpine Research.

His research focuses on plant ecology, biogeochemistry, and community dynamics.

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作者 III ： Sally D. Hacker

Received her Ph.D. from the Department of Ecology ad Evolutionary Biology at Brown University.

Associate professor in the department of Zoology at Oregon State University.

Research explores species interactions and how they influence community formation, species diversity, and species invasions.

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Preface (01) This is an exciting and challenging time to

study ecology. 應用於面對與化解環境問題 新技術的運用，分子技術，衛星影像技術 熱門的學科 學生需要學習多元多樣的知識和能力 缺少適宜的 ( 好的 ) 教科書

Two core principles that guided our writing. Teaching comes First. Less is More.

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Preface (02) Features

1. Pedagogical excellence2. Subject matter3. Case studies4. Connections in Nature5. Ecological inquiry6. Hands-on Problem solving7. Ecological applications8. Links to evolution9. Art Program

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Supplements Companion website

教科書的網站： http://www.sinauer.com/ecology

Ayo NUTN website： ( 教學網站 ) http://

myweb.nutn.edu.tw/~hycheng/

T01. 簡介：生態學Chap.01 The Web of Life

鄭先祐 (Ayo) 教授國立台南大學 環境與生態學院

生態科學與技術學系 環境生態研究所 + 生態旅遊研究所

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Chap. 1 Introduction: The Web of Life

Case Study: Deformity and Decline in Amphibian Populations

Introduction 1.1 Connections in Nature1.2 Ecology1.3 Answering Ecological QuestionsCase Study RevisitedConnections in Nature: Mission

Impossible?

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Case Study: Deformity and Decline in Amphibian Populations

Figure 1.1 Deformed Leopard Frogs

High incidence of deformities in amphibians

Declining populations of amphibians worldwide

Many declining populations were in pristine or protected areas.

Amphibians are “biological indicators” of environmental problems.

Air and water pollution, changes in temperature and in the amount of ultraviolet light.

Ayo 2011 Ecology18Figure 1.2 Amphibians in Decline

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Introduction

Humans have enormous impact on the planet.

We must understand how natural systems work.

Ecology: The scientific study of how organisms affect—and are affected by—other organisms and their environment.

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1.1 Connections in Nature

Concept 1.1: Events in the natural world are interconnected.

Even species that do not interact directly can be connected by shared environmental features.

Ecologists ask questions about the natural world to understand these connections.

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Observation and test hypothesis

觀察： Observation of the deformities of pacific tree frogs and long-toed salamanders. 証據： the deformed amphibians all contained a

parasite, known to be Ribeiroia ondatrae, a trematode ( 吸蟲類 ) flatworm ( 扁形蠕蟲門 ).

Ruth 提出假說： hypothesized that a parasite can cause deformities. 驗證： Implanted glass beads( 玻璃小珠 ) mimic

the effect of the cysts ( 囊胞 ) of the parasite. 結果： The beads caused deformities similar to

the parasite.

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Johnson’s work (01)

35 ponds in Santa Clara county.13 ponds with Pacific tree frogs,

4 of 13 ponds contained deformed frogs. 2 of 4 ponds ，高達 15-45% of tadpoles in

metamorphosis 有畸形。 原本以為是水污染的結果。 Pesticides,

PCBs, or heavy metals 等，但於這 2 ponds的水中都沒有發現。

採集 200 顆卵回實驗室，成長都正常。因此並不是遺傳。

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Johnson’s work (02)

The 4 ponds with abnormal frogs were the only ponds that contained both tree frogs and an aquatic snail, Planorbella tenuis. The snail is an intermediate host in the life

cycle of the Ribeiroia parasite (Fig. 1.3) Ribeiroia cysts in all the frogs with

deformed limbs.

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Figure 1.3 The Life Cycle of Ribeiroia

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Johnson’s work (03)

假說： Ribeiroia caused deformities in Pacific tree frogs.

驗證： controlled experiment Tree frog eggs were exposed to Ribeiroia

parasites in the lab. Four treatments: 0 (the control group), 16,

32, or 48 Ribeiroia parasites. 結果 (Fig. 1.4)

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Figure 1.4 Parasites Can Cause Amphibian Deformities

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A field experiment: Kiesecker (2002)

現象： Six ponds, all of with contained Ribeiroia,

but only some of which contained pesticides Six ponds, three with pesticide

contamination.野外實驗方法： (Fig. 1.4)

Six cages in each pond, three with mesh size that allowed parasite to enter, the other three had a mesh too small for the parasites.. (Fig. 1.5a)

結果 (Fig. 1.5b)

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Figure 1.5a. Do Ribeiroia and Pesticides Interact in Nature?

Ayo 2011 Ecology29Figure 1.5b, Do Ribeiroia and Pesticides Interact in Nature?

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1.1 Connections in Nature

Hypothesis: Pesticides decrease the ability of frogs to resist infection by parasites.

Lab experiment: Tadpoles reared in presence of pesticides had fewer white blood cells (indicating a suppressed immune system) and a higher rate of Ribeiroia cyst formation. (Fig. 1.6)

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Figure 1.6 Pesticides May Weaken Tadpole Immune Systems (Part 1)

Tadpoles exposed to the pesticide had fewer eosinophils

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Figure 1.6 Pesticides May Weaken Tadpole Immune Systems (Part 2)

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1.1 Connections in Nature

Synthetic pesticide use began in 1930s; use has increased dramatically.

Amphibian exposure to pesticides has also increased.

Any action (increased pesticide use by people) can have unanticipated side effects (more frequent deformities in amphibians).

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1.1 Connections in Nature

Fertilizer use may also be a factor:Fertilizer in runoff to ponds increases

algal growth.Snails that harbor Ribeiroia parasites

eat algae.Greater numbers of snails result in

greater numbers of Ribeiroia parasites.

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1.1 Connections in Nature

Many human actions have also increased human health risks.

Damming rivers in Africa increases habitat for snails that carry schistosomiasis.

New diseases, such as AIDS, Lyme disease, Hantavirus, Ebola, and West Nile fever (Fig. 1.7) may be related to human actions.

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Figure 1.7 Rapid Spread of a Deadly Disease (Part 1)

West Nile Fever

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Figure 1.7 Rapid Spread of a Deadly Disease (Part 2)

West Nile Fever

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1.2 Ecology

Ecology is a branch of biology.Environmental science incorporates

concepts from the natural sciences (including ecology) and the social sciences, and focuses on solutions to environmental problems.

Concept 1.2: Ecology is the scientific study of interactions between organisms and their environment.

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1.2 EcologyEarly ecological views:1. There is a “balance of nature,” in which

natural systems are stable and tend to return to an original state after disturbance.

2. Each species has a distinct role to play in maintaining that balance.

Scientists now recognize that ecological interactions are more complex.

One view that stood the test of time: Events in nature are interconnected.

“You can never do just one thing.”

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1.2 EcologyEcologists study interactions in nature

across many levels of organization.Ecological studies usually emphasize

individuals, populations, communities, or ecosystems.

Levels of Biological organization (Figure 1.8)

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Figure 1.8 Levels of Biological Organization (Part 1)

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Figure 1.8 Levels of Biological Organization (Part 2)

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1.2 EcologyA population: A group of individuals of a single

species that live in a particular area and interact with one another.

A community: An association of populations of different species living in the same area.

An ecosystem: A community of organisms plus the physical environment in which they live.

All the world’s ecosystems comprise the biosphere — all living organisms on Earth plus the environments in which they live.

Ayo 2011 Ecology47Figure 1.9 A Few of Earth’s Many Communities

(A) Savanna(B) Rainforest

(C)Dunes in the Namib desert

(D)Shallow-water marine community

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Ecology is broad in scope and scale

Every ecological study must be done at an appropriate scale, both spatially and temporally.

Small spatial scale: Soil microorganisms.

Large spatial scale: Atmospheric pollutants.

Short temporal scale: Leaf response to sunlight.

Long temporal scale: How species change over geologic time.

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Key terms (Table 1.2)

All living systems change over time.

Evolution:1. A change in the genetic characteristics

of a population over time.2. Descent with modification—organisms

gradually accumulate differences from their ancestors.

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Key terms

Natural selection: Individuals with particular adaptations tend to survive and reproduce at a higher rate than other individuals.

If the adaptation is heritable, the offspring will tend to have the same characteristics that gave their parents an advantage.

As a result, the frequency of those characteristics may increase in a population over time.

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Figure 1.10 Natural Selection in Action

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Key terms

Ecosystem processes: Movement of energy and materials.

Energy enters the community when producers capture energy from an external source, such as the sun, and uses that energy to produce food.

Net primary productivity (NPP): Energy that producers capture by photosynthesis or other means, minus the amount they lose as heat in cellular respiration.

Energy moves through ecosystems in a single direction only—it cannot be recycled.

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Figure 1.11 How Ecosystems Work

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Key terms

Nutrients are continuously recycled from the physical environment to organisms and back again.

Nutrient cycle: Cyclic movement of nutrients such as nitrogen or phosphorus between organisms and the physical environment.

Life would cease if nutrients were not recycled.

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1.3 Answering Ecological QuestionsConcept 1.3: Ecologists evaluate competing hypotheses about natural systems with experiments, observations, and models.Ecologists use several methods to answer

questions about the natural world:

1. Observational studies in the field.

2. Controlled experiments in the laboratory.

3. Experiments in the field.

4. Quantitative models.

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1.3 Answering Ecological Questions

An observational field study: Johnson et al. (1999) surveyed ponds to determine

that frogs with deformities were only present if the parasite’s intermediate host snail was also present.

A controlled experiment: Johnson et al. (1999) also tested their observations

by exposing tadpoles to different levels of the parasite Ribeiroia in the laboratory.

A field experiment: Kiesecker (2002) compared frogs from three ponds

containing pesticides with frogs from three ponds that had no pesticides.

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Table 2.3

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The scales do matter

Ecological experiments can be done at different scales:

Small-scale laboratory experiments in test tubes or flasks, to whole-lake experiments.

Sometimes experiments are difficult or impossible to perform. Example: When questions concern events

occurring over large geographic scales, such as global warming.

The study of global warming involves using a mixture of observational studies, small-scale experiments, and quantitative (mathematical or computer) models.

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Figure 1.12 Ecological Experiments

(A) Laboratory experiments

(B) small-scale field experiments

(C)Large-scale experiments

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Experimental design:

1. Replicate—perform each treatment more than once.

2. Assign treatments at random.3. Statistical analysis is used to determine

significant effects.

Replication: As the number of replicates increases, it becomes less likely that the results were actually due to a variable that was not measured or controlled.

Assigning treatments at random helps to limit the effects of unmeasured variables.

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Figure 1.13 Experimental Design and Analysis

This study was designed to test the effects of insect herbivores on plant growth and asexual reproduction.

A set of 5x5 meter plots.

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Scientific method:

1. Make observations and ask questions.2. Use previous knowledge or intuition to

develop possible answers (hypotheses).3. Evaluate hypotheses by performing

experiments, doing observational studies, or using quantitative models.

4. Use the results to modify the hypotheses, to pose new questions, or to draw conclusions about the natural world.

5. The process is iterative ( 反覆的 ) and self-correcting ( 自我修正的 ).

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Case Study Revisited: Deformity and Decline in Amphibian Populations

Studies have suggested that no single factor can explain decline of amphibian populations. The declines seem to be caused by complex

factors that often act together and may vary from place to place.

Hatch and Blaustein (2003) studied the effects of UV light and nitrate on Pacific tree frog tadpoles. At high elevation sites, neither factor alone had

any affect. But together, the two factors reduced tadpole survival.

At low elevation sites, this effect was not seen.

Ayo 2011 Ecology65Figure 1.14 Joint Effects of Nitrate and UV Light on Tadpole Survival

Joint effects

Synergetic effects

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Case Study Revisited: Deformity and Decline in Amphibian Populations

The effects of pesticides are also complex. Some studies show that tadpoles are

more susceptible when under stress, such as presence of predators.

A broad set of factors can cause frog deformities, but little is known about how these factors interact. There remains much to be discovered in

the field of ecology.

Ayo 2011 Ecology67Figure 1.15 Complex Causation of Amphibian Deformities

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Connections in Nature: Mission Impossible?

The natural world is vast, complex, and interconnected.

But ecologists think that it is not impossible to understand it. Ongoing efforts are sure to be

challenging, exciting, and important to the well-being of human societies.

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科學研究者的懷疑論

Scientists demand hard evidence.Most good scientists are skepticsAny idea, hypothesis, or experiment is

certain to be challenged by someone.Like hypotheses, theories are open to

tests, revision, and tentative acceptance or rejection.

補充資料

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科學驗證的可能錯誤

型一錯誤 (Type I error) 否定 null 假說，但事實是「 null 假說是成立的」 the conclusion that there is an effect when in fact

there is none, ( 否定 null 假說，但事實是「假說是成立的」 )

型二錯誤 (Type II error) 接受 null 假說，但事實是「 null 假說是錯誤的」 when an impact exists but is not detected. ( 接受

null 假說，但事實是「假說是錯誤的」 )

補充資料

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型一 (Type I) 錯誤 vs. 型二 (Type II) 錯誤

驗證結果 真實情況

拒絕 Ho 接受 Ho

Ho是真實 結果錯誤

(型一錯誤) 結果正確

Ho是錯誤 結果正確 結果錯誤

(型二錯誤)

補充資料

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下降錯誤的目的與取樣方法：型一 vs. 型二

下降型一錯誤： 目的：儘可能的不要 拒絕「零假說」。 取樣：隨機，擴大範圍與數量

下降型二錯誤： 目的：盡可能的不要 接受「零假說」。 取樣：重點，按最可能地點 ( 重點 ) 取樣。

補充資料

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自然科學 vs. 公害的研究

自然現象的研究，盡量的下降 型一錯誤。 盡量避免否定「零假說」。 如此可以避免研究方向的錯誤。

對於公害或生態保育的研究，則需要選擇下降 型二錯誤。 盡量要否定「零假說」。 公害往往隱藏，受害往往有空窗期。警訊出

現，往往如同冰山一角。 倘若遺漏，等到其爆發，災難難以承擔，或已經難以挽回。

補充資料

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自然運作，廣泛。 取樣宜擴大範圍，

隨機取樣人為運作，集中。

取樣若擴大範圍，隨機取樣，遺漏重點的機率增加。

盡量避免萬一沒有發現問題

補充資料

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Dioxin 污染調查

案例：台江國家公園的戴奧辛污染調查。如何取樣？

下降「型一錯誤」的方法• 平均隨機取樣，廣泛取樣。• 未必可找到污染地點。

下降「型二錯誤」的方法• 找尋最有可能受污染的地點，密集取樣• 務必要找到污染地點。

補充資料

生態學定位與發展史

鄭先祐 (Ayo) 教授國立台南大學 環境與生態學院

生態科學與技術學系 環境生態研究所 + 生態旅遊研究所

補充資料

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「生態學」的定位 (起源 )

第一個使用 ecology (Haeckel, E., 1866) 。 Ecology as a part of physiology ( 生理學 ) 。

名詞： Ecology 字源取自 eco-nomics (household+management) eco-logy (the study of household) the economy of nautre ( 自然的經濟 ) 。

內容： 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.

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生態學 與 經濟學Eco-logy and Eco-nomics

都是 Eco ( 生活 ) 之學。 Eco + logy = 生活 + 科學 Eco + nomics = 生活 + 經營管理。

生態學 = 「自然的經濟」(Economy of Nature)

什麼才是真正的「經濟」？

3

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生態學 與 經濟學

海洋生態體系

物質循環

能量循流

太陽

捕魚活動

人力、漁具、漁船、能量

海產市場

經濟學 生態學

圖 8. 海灣區的經濟活動。左半部是傳統生態學的研究領域，右半部是傳統經濟學的課題。海灣區對人類的實際價值至少是此區生產的魚獲量價值的 10倍。

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經濟學 vs. 生態學

Odum, E. P. and G. W. Barrett (2005) Fundamentals of Ecology.

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「生態學」的定位 ( by 1910s)

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)

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「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 Kuhn (1970) a developing scientific

discipline ==> fusion of several separate trunks lacking a common initial rootstock.

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「生態學」的定位 ( 1960-80s)

1961 Andrewartha 生態學 = 生物的分佈和豐富量之研究。 1963 Odum, E.

生態學 = 自然的結構和功能之研究。 1980s Odum, E. (1983) 生態學 = 自然科學和社會科學的橋樑。

Leo Smith, R. (1986) 生態學 = 「生物與其生活環境」的互動關係之研究。

生態學 = 自然的經濟學

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「生態學」的定位 ( 1990s)

Stiling, P.D. (1992) 生態學包含以下五部份：

演化生態 + 行為生態 + 族群生態 + 群落生態 + 應用生態

Leo Smith, R. (1992) 生態學 = 主要兩個根源：植物地理學 和 自然歷史學 Colinvaux, P. (1993)

生態學 =the Science of the universe• 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

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「生態學」的定位 ( 1990s)

Odum, E. (1993) 生態學 = 生命與維生環境之關係。

Krebs, C. J. (1994) 生態學 = 決定「生物的分佈和豐富量」之互動的研究。

Where, How many, WhyBrewer, R. (1994)

生態學 =

「生物與其生活環境」的互動關係之研究。

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「生態學」的定位 ( by 2000)

Stiling, P. D. (1996) 生態學 = 生物和生物與其生活環境間的互動關係。Ricklefs, R. E. (1997)

生態學 = 自然的經濟。Smith and Smith (1998)

生態學 = 自然的經濟。Ricklefs and Miller (2000)

生態學 = 研習 自然世界的 次序 (order) 。

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生態學發展史 (一 )

1970s Conservation + Landscape + Restoration

1960s Ecology 逐漸被認為是重要「科學」。 1950s 各自發展屬於自己領域的「人類生態學」。 1940s

1930s

1920s 生態學的理論基礎 1910s Animal Ecology + Human Ecology

1900s Ecology 成為「生物學」中三大部之一。 1866 Haeckel, Ernst : 第一個使用 Ecology

Social ecology + Cultural ecology

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2000s Business Ecology + Information Ecology + Knowledge Ecology + 大學系所(doctoral degree)

1990s Journal of Industrial Ecology + Eco-efficiency ( 生態效率 )

1980s Eco-Industrial park + Deep Ecology + Spiritual Ecology + Radical ecology + Eco-ethics

1970s Industrial Ecology + Conservation + Landscape + Restoration + EIA

1960s Ecology 才逐漸被認為是重要的「科學」。

生態學發展史 (二 )

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現代生態學的類別 生物 : Bird Ecology, Lizard Ecology, Plant

Ecology, Fish Ecology 演化 : Population Genetics, Evolutionary Ecology 理論 : Theoretical Ecology, System Ecology,

Ecosystem Ecology, Physiological Ecology, Behavioral Ecology

環境 : Environmental Conservation, Conservation Biology, Landscape Ecology, Geographical Ecology

社會 : Social Ecology, Human Ecology, Ecological Impact Assessment,

事業： Business Ecology, Industrial Ecology, Information Ecology

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多元途徑： 人類生態學 (Human Ecology) 生態經濟學 (Ecological Economics) 工業生態學 (Industrial ecology) 事業生態學 (Business Ecology) 資訊生態學 (Information Ecology) 藍海策略 (Blue ocean strategy) 美學的經濟 (the economy of Aesthetics) 生物模擬 (仿生學 ) (Biomimicry) 自然之道 (The Natural Step)

新世紀的生態學

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學術發展：智價革命人類生態學：自然與人文的對話、互動與整合復育生態學：都會區生活圈的物種復育事業生態學：事業生態價值化工業生態學：工業生態程序化生物模擬 (Bio-mimicry) ：向自然學習

生態科學與技術的發展

Ayo NUTN website:http://myweb.nutn.edu.tw/~hycheng/

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