PAGE PROOF: 2ND PASS

C H A P T E R

1 The Science of Plant Ecology

T
he biological science of ecology is the study of the relationships between
living organisms and their environments, the interactions of organisms
with one another, and the patterns and causes of the abundance and dis-
tribution of organisms in nature. In this book we consider ecology from the
perspective of terrestrial plants. Plant ecology is both a subset of the discipline
of ecology and a mirror for the entire field. In The Ecology of Plants, we cover
some of the same topics that you might find in a general ecology textbook,
while concentrating on the interactions between plants and their environments
over a range of scales. We also include some subjects that are unique to plants,
such as photosynthesis and the ecology of plant-soil interactions, and others
that have unique aspects in the case of plants, such as the acquisition of
resources and mates.

Ecology as a Science
Ecologists study the function of organisms in nature and the systems they
inhabit. Some ecologists are concerned in particular with the application of
ecological principles to practical environmental problems. Sometimes the
distinction between basic and applied ecology becomes blurred, as when the
solution to a particular applied problem reveals fundamental knowledge about
ecological systems. In both basic and applied ecology, the rules and protocols
of the sciences must be rigorously followed.
What ecology is not is environmental or political activism, although ecol-
ogists are sometimes environmental activists in their personal lives, and envi-
ronmental activists may rely on ecological research. Ecology is not about one’s
feelings about nature, although ecologists may have strong feelings about what
they study. Ecological systems are complex things, with a great many parts,
each of which contributes to the whole in different ways. Nevertheless, ecolo-
gy is indeed a science, and it works like other scientific disciplines.
How do we know whether something is true? Science is one way of know-
ing about the world—not the only way, but a spectacularly successful one. In
contrast to some of the other ways of knowing that are part of our lives, the
legitimacy of science is based not on authority, or opinion, or democratic prin-
ciples, but on the weight of credible, repeatable evidence.

(Table 1. and Einstein’s theory of relativity. They carry out a series of steps. is built on a tri- pod of pattern. or even a guess or “hunch. known before. using assumptions. ecologists gather various kinds of information. followed by retest. these science than it does in common usage. Eventually. Speculate. but out of this chaos ory of evolution. hypothesis. and verification. comes our understanding of nature. among other things. Formulate hypothesis sist of the relationships between pieces or enti. it is essential: every scientific study must begin by assessing what is already known. over time. Processes are the Reevaluate observations causes of those patterns. When a theory is buttressed over many years with struct theories. But the heart Figure 1. The cycle of speculation. and the strong evidence. (often phrased as a question) ties of the natural world. Scientists use such overarching theories to organize . Darwin’s the- be a messy and chaotic process. Although there are exceptions. That experience of discovery is what makes doing science so incredibly exciting and fun. Science in operation can occurred with Newton’s theory of gravity. and theory nations of those causes. the word “theory” usual- ing of the new or modified hypotheses. process. mischief in the popular press and in public debates on terns. and ultimately to put together theories that More predictions. The scientific method. In ecology. Compare dation based on an enormous collective store. rejec. like all of science. When ecologists carry Predict results assuming out original scientific research. although not The word “theory” has a very different meaning in always in a fixed order (Figure 1. comprehensive explanation of a large description. explain what they have found out.2 Chapter 1 PAGE PROOF: 2ND PASS The Genesis of Scientific Knowledge Observations Throughout this book we examine how ecolo. in advancing their understanding. data. they con. Patterns con. with new findings consistently sup- results of many tests of hypotheses. must be support- those hypotheses using experiments (in a broad sense of ed and ultimately confirmed (or rejected) by the accu- the word. This sort of research is not only useful. They often put together some sort of model to help politically charged issues. and data gists have come to their current knowledge and understanding of organisms and systems in nature. Apply inductive oretical basis and has developed from a foun. research carried out by students or the general public is usually secondary research: Independent verification gathering data or confirming facts that are of results by other researchers: “reproducible results” already known. scientific theory with “a guess” has caused no end of pose processes that might be responsible for those pat. testing body of information that. Ecology has both a strong and a rich the. with current theories. ory is a broad. posing hypotheses. Design experiment(s) to test further development validity of predicted results There is a distinction between the kind of theory of research a scientist does and the kind of research done for a term paper. This is what has sometimes at cross-purposes. and theory.1). Scientists gain knowledge by using the scientific method. understand processes. sometimes working in synchrony and which new speculation can spring.” Equating the meaning of a look for patterns or regularities in their data and pro. Throughout this ly refers to a limited. and experimentation is circular. Ecology. mulation of a wide range of different kinds of evidence tion. A scientific the- steps can be summarized as follows: observation. Theories are the expla. porting and amplifying the theory while producing no The building of comprehensive scientific theories pro. or revision of the hypotheses.1). or by Results support Results do not any member of the public trying to gather hypothesis (predictions support hypothesis information about a topic using library books or confirmed) (“null hypothesis”) material posted on Web sites. and deductive reasoning to observations. further experiments. quantification. they seek to hypothesis to be correct document patterns. process. specific conjecture or supposition. models. serious contradictory evidence. as new questions constantly ering information or finding out facts that no one has ever emerge from the answers scientists obtain.1 of what research scientists do is primary research: gath. it becomes an accept- ceeds simultaneously from multiple directions by ed framework or pattern of scientific thought from numerous people. as discussed below). In popular usage. house of information about natural history.

made of plastic order to reach a genuinely new understanding. is a species’ Assumptions Conditions or structures needed to build the theory population size so small that it Concepts Labeled regularities in phenomena is becoming endangered? How Definitions Conventions and prescriptions necessary for the theory to work rapidly is an invasive species with clarity spreading? How many species Facts Confirmable records of phenomena can coexist in a community. examining the weight els are used to define patterns. civil engineers often build discoveries. A crucial characteristic of science is the need to col. Build. While one tions. designed for efficiently conveying essential infor- revise or reject a hypothesis if the evidence does not sup. must be objective in. The justifications for the research presented in a attempts to understand complex structures and rela. in Scientific Research entific evidence. What one chooses to study. it may at first we discuss their design and use in more detail later in this seem arcane and dull. tivity and creativity must also come into play. and because faulty assumptions and unjus- A scientific hypothesis is a possible explanation for tified simplifications can sink even the most widely a particular observation or set of observations. Subjectivity. and Chance tion or statement that can be verified or rejected using sci. Mod. We will let you in fication that expresses structures or relationships. especially in ecology.1 The components of a theory rely on mathematics is that ecologists are so often con- Component Description cerned with the numbers of Domain The scope in space. While everyone knows that science which we will discuss at length in Chapter 22. laboratory or field disasters. Hypotheses must be testable: they must contain a predic. Mathe- Laws Conditional statements of relationship or causation. time. or even ing a model airplane from a kit can tell you a lot about after the work was finished. Ecologists deal almost research. plicit) simplifications and assumptions are critical to their thinking and derive additional predictions about recognize because they can alert you to the limitations nature. The format follows a rigid proto- chapter. with a clear logical line running from start to finish. would find even before they began. Because of serendipitous the basic form of an airplane. come from a and you may have heard about global climate models. whether in science or in everyday life. PAGE PROOF: 2ND PASS The Science of Plant Ecology 3 One reason the models often Table 1. In science. esis is smaller in scope than a fully developed theory. and what is or is not a valid topic simple verbal argument to a set of mathematical equations. Choice. is objective and rational. for example. the original purpose of a research proj- (either physical models or three-dimensional images on ect is sometimes modified or. and When you read a typical scientific paper. sets of equa. Ideas in science. where one choos- exclusively with abstract models that can range from a es to look for answers. One of the most valuable uses of a subtle but important role throughout all of scientific models is to make predictions. Science does not accept hypotheses on faith. or statements matical models offer well-de- of process that hold within a universe of discourse fined methods for addressing Models Conceptual constructs that represent or simplify the natural world questions in both qualitative Translation modes Procedures and concepts needed to move from the abstractions and quantitative terms. that is only half the story. occasionally. In Models can be abstract or tangible. of a theory to the specifics of application or test All models are necessari- Hypotheses Testable statements derived from or representing various components of the theory ly based on simplifications Framework Nested causal or logical structure of the theory and rest on a set of assump- tions. Those (implicit and ex- Source: Pickett et al. subjec- or words. It Some of the most important tools in the scientist’s may seem as if the researchers knew exactly what they toolkit are models. Confirmed generalizations Condensations and abstractions from a body of facts that have and how does this change as been tested conditions change?). summarize processes. for scientific research are all subjective decisions scien- . variety of sources. covered long after the research project began. subjectivity plays generate hypotheses. 1994. and of evidence in support of a hypothesis. and phenomena addressed by the theory things (for instance. accepted or elegant model. entirely dis- a computer) before construction is begun. on an open secret: That is not usually how real science els are one of the ways in which the human mind works. Objectivity. or unusual nat- small models of structures such as bridges or buildings ural occurrences. Ideas are tightly packaged. mod. A model is an abstraction or simpli. A hypoth. Experiments are the heart of science. or a complex computer program. paper’s introduction may have been thought up or dis- tionships. They can be diagrams on paper. doubt seen models of DNA and of chemical reactions. You have no carded and replaced with something else. port it. of the model. mation to other scientists.

community. den. The controlled environment large scales of space and time—for example. however. or in a natural community in a field setting. ses. sacrificing generality. the early twentieth century. and tries again. If the experiment is con. Or an idea the social sciences. Ecology is con- working in a controlled environment. Such ation so that the differences among treatments can be experiments have limitations. but now many factors ing the answers must be objective. Coming up with a good experiment. By Another limitation is that of scale. Ecological experiments “poke-at-it-and-see-what-happens” experiments designed can be classified into three broad types: manipulative. because she can separate factors that gist is able to control most of the possible sources of vari. common throughout the biological sciences. and thus their works in a particular way? Or a previous experiment results may be more readily generalized. is largely subjective. the scientist gains confidence that his hypothesis Ecologists in particular use a wide variety of types of might be correct. and second. cellular biology as well as the physical sciences. ment. this continuum of control versus realism the ecologist tist successful is the ability to recognize the worth of carries out her experiment depends on both her scien- these casual observations. and new tific goals and practical considerations. Good experiments avoid artifacts experiments are the mainstay of most of molecular and or take them into account in evaluating the results. parts of the natural world will be altered to study their ducted in a laboratory or growth chamber. and measure such things as these general types of experiments. Fisher in creative activities. he is guid- their time to maturity and their final size. From these sources. Baconian treatment being tested. is why each study expands our ecological knowledge. Manipulative experiments are powerful tools for two This range of potential settings for the experiment major reasons: first. and con- switching gears after a disastrous laboratory failure to clusions are based on the use of statistical inference (see extract a successful outcome from the jaws of catastro. more realistic or more natural. in an experimental gar. questions. nat. or the responses of populations to climate ited range of conditions). looking at a ation. what-if thoughts. Instead of attempting to control all vari- well. revises his ideas. Fisherian experiments are a Many scientific discoveries start with casual obser. effects. This sort together to build a comprehensive theory are all highly of experiment was first developed by R. the Appendix). We cannot do . may vary in an uncontrolled fashion. cerned with patterns and processes that occur across ogist sacrifices something. choosing what ques. ment. and pulling a large number of disparate facts many settings and are not restricted to the field. Arabidopsis thaliana. however. Manipulative.4 Chapter 1 PAGE PROOF: 2ND PASS tists must routinely make. typically occur together to test them individually. and observational. Scientists have studied in detail only a few hun- A person manipulates the world in some way and looks dred of the quarter-million terrestrial plant species. such as Newton’s proverbial apple. experiments are experiments. They are typically less narrowly may arise as a “what if” thought: What if the world defined than Baconian experiments. the sci- entist goes back. mental ones is randomized among replicates. and how to ask them. including experiments are what most of us think of as experiments: ecology. In a field experi- tions to ask. in a greenhouse. This experiment ed by his knowledge of other similar species and ecosys- could be done in a controlled environment such as a tems. the causes is highly artificial so that it compromises realism. If the Experiments: The Heart of Research hypothesis is not falsified by the outcome of the experi- A cornerstone of the scientific process is the experiment. which growth chamber. however. If an experiment is conducted in a field setting. or controlled. or ecosystem must do many of different nutrient treatment. For example. the ecol. While determin. variation due to factors other than the experi- seemingly intractable problem from a new perspective. it is es depend on the questions one asks. the only factors that are controlled are the ones Many scientific endeavors are highly creative as being studied. only a few [Zea mays (maize). These are broadest sense: a test of an idea. Of course. If the hypothesis is partially or wholly falsified. Such experiments can be carried out in phe. an ecologist constructs Experiments are usually designed as tests of hypothe- hypotheses and designs experiments to test them. (rice)] approach being “thor- nutrients on the growth of a particular plant species. She oughly studied. and of differences in the numbers of species on different con- it is also narrow in scope (the results apply only to a lim. We use the term “experiment” here in its done without an explicit hypothesis in mind. because the scientist can control which comes with a set of trade-offs. an ecologist these. giving each of them a new species. these choic. Each species or ecosystem is unique. A. change over the next two centuries. the ecolo. To a large extent. to find out more about the world. Such experiments are ural. however. Sometimes. tinents. of for a pattern in the response. Where along may have raised new questions.” An ecologist beginning the study of a can grow several groups of plants. caused by a side effect of the experimental manipulation plify the scientific method as it was first laid out by itself rather than being a response to the experimental Frances Bacon in the seventeenth century. Sometimes manip- clearly attributed to the factors being studied in the ulative experiments are plagued by artifacts—outcomes experiment. might be interested in the effects of different amounts of and possibly Oryza spp. mainstay of ecology and evolutionary biology as well as vations. What makes a scien. These sorts of controlled experiments exem.

which are watershed at the Konza Prairie Research Natural Area (A). 3. One example of such a study is the Large-scale manipulative experiments are often lim- long-term study of prairie ecology at the Konza Prairie ited by the range of possible treatment. F. and S. Because we do not . B. H = Replicate indicators 4A 4B 10A 20B SuB 4F 20A Season of burn treatment: WA WH 4H SpB FH W. 1998. In burns (B) are done at various intervals to investigate the this map. Konza Prairie Biological Station. vary in size from approximately 3 to 200 hectares. which are subjected to dif- and space. Summer. Fall 1C 2B 10B FA Figure 1. 1998). In fire treatment.2 pared with ungrazed areas and with plots subjected to cattle Large-scale manipulative experiments are being carried out grazing. 20 = Number 2B of years between 4D 1B burns 1D 2D 4A 4D 20D A. however. 2. The reserve is divided burning is carried out in the spring. PAGE PROOF: 2ND PASS The Science of Plant Ecology 5 (A) (B) (C) (D) N 0 1 2 3 km AL 3C AL 3A Ungrazed 1A A HQ HQB 20A 3B Grazed by bison AL HQC Grazed by cattle 4A 2A 20A 4A Agricultural land 1A 4B 4B 2A 1B HQ Headquarters 1B area (small 20B experimental plots) 4C 1C 1. 4. each patch is designated by a code indicating the effects of fire and fire frequency on prairie communities. at Konza Prairie. areas grazed by bison (C) are studied and com. of A. (After Knapp et al. begun by Lloyd Hul. example. iments (see Box 12C). 2A SpA 20C Sp. Knapp. 1B 2C 1A SuA Su. time intervals and grazing by bison or cattle (Figure 1. Photographs courtesy addition. for Research Natural Area in Kansas. increasingly making ferent combinations of controlled burning at various use of longer-term and larger-scale manipulative exper. however. 10.) manipulative experiments at these great scales of time into a series of large patches.2). Ecologists are. Collins. C. almost all of the controlled bert in 1981 (Knapp et al. D. Winter. Prescribed units. The experimental patches (D). Spring.

then she gains confidence about the This assumption goes against our own experience and causes of those changes. it is so on. If an ecologist finds similar thing to be true.3). ones. ductivity.6 Chapter 1 PAGE PROOF: 2ND PASS have firm data on prairie fire regimes prior to European ecologist cannot tell for sure whether the increase in her- settlement. it can be First. similar to the trade-off lenging and exciting science. might have had a differ. to falsify the hypothesis. philosopher of science Karl Popper (1959). In such cases. all the- The best natural experiments are ones that repeat ories are held to be potentially false. These are natural and observational gy and astronomy. a volcanic eruption may denude mation gained from many different sources and many an area. There are hypotheses. even if we still do not know the details of that Observational experiments consist of the system. if the al times in a row. if we are comparing areas burned in a major hypothesis or theory. is a more subtle. the outcome of the test either falsifies or fails areas might have been wetter. standing. we propose a hypothesis fire with others that remained unburned. Bang. many facts that the chance that we are wrong is very lated. The popular image of there is never just a single difference before and after a the scientific method portrays it as a process of falsifying change or between systems being compared. We know that life began and assumed its pres- atic study of natural variation. Rather. In this frame- tered systems were identical prior to the event. not by “humours. This approach was codified by the German no guarantees. nuanced. this approach fails to recognize knowledge difficult to determine the cause of any change. unethical to carry out. Natur. we do not know how this spring-burning bivores is a result of increased plant numbers and pro- treatment compares to “natural” fire regimes. and falsification is an important part of theory testing. but has only productivity (see Chapter 20). While hypothesis testing and ent site history or different vegetation before the fire. In a strict Popperian framework. there are many other potential not the whole story. In other words. the of knowledge (Mayo 1996). The ecologist’s use of this al and manipulative experiments represent a trade-off complex variety of information makes ecology a chal- between realism and precision. the unburned and test it. the ecologist compares the Testing Theories altered system either with the same system before the The testing of scientific theories. notably geolo- types of studies. Other sciences. For example. sources of difference besides the fire. The school of philosophy of science number of herbivores is also observed to increase as the called realism recognizes this progressive accumulation number and productivity of plant species increases. Today we know that the Earth revolves around ments. For work. event. While we may acknowl- across a continent to test hypotheses about the relation. one could measure patterns of species diversity veyed by DNA. or whether the increased productivity is a Some types of manipulative experiments would be result of increased herbivory. a species may Ecological knowledge comes from combining infor- go extinct in a region. If several factors are tightly corre. We know measurements are experiments if an ecologist starts with that many diseases are caused by microbial infections. We never prove any- themselves in space or time. which may be thought of as different kinds of observational experiments because of the spatial or tem- experiments. Just as with a manipulative experiment. . we would not cause As with natural experiments. changes each time. accumulation. For exam. not by blood.” and that hereditary traits are con- ple. Another approach is to com. for two reasons. edge that all of this knowledge has not. that the altered and unal. we merely disprove ideas that are false. Such observations or ent shape through the process of evolution. For example. in a strictly ship between the number of plant species number and philosophical sense. been proved to be true. and complicated endeav- The major limitation of natural experiments is that or than nonscientists often realize. by some natural occurrence. For example. for instance. also rely largely or exclusively on studies. old (give or take a few billion) and began with the Big imental manipulation. the history of the accumulation of scientific under- bine natural experiments with manipulative experi. unchanged system. the patches subjected to grazing the sun. knowledge is so firmly established and bolstered by so tors to vary together. observational experi- the extinction of a species just to study the effects of such ments repeated in space or time add confidence to our an event. it becomes difficult to determine which factor is much less than the chance of winning the lottery sever- the cause of the observed pattern. Again. poral scales of their studies or because direct manipu- Natural experiments are “manipulations” caused lation is impossible. or a flash flood may scour a streambed. We and fire treatments at Konza Prairie are being compared know that the universe is approximately 15 billion years with patches elsewhere that are not subjected to exper. For example. one or more hypotheses (predictions) to test. different kinds of experiments. we are taught that we can never prove a scientific example. especially ecological change or with a similar. even though this was once just a hypothesis. the limitation of failed thus far to be falsified. between laboratory and field experiments. ecologists must rely on two other conclusions (Figure 1. we also recognize that some this type of experiment is the potential for multiple fac. Therefore.

dif- 130 ferent interpretations of experimental data. we ask. the Popperian frame. but expanding the context with increasing numbers 90 90 of observations over time shows that (B) 1998 was the warmest winter in 10 years. “How much. Ultimately. judgment. then the matter becomes settled (unless ask. we do not of its validity. yes or no. But when the evidence in favor of a scientific one of falsifying a hypothesis. records of the duration of ice cover on Lake Duration of ice covert (days) Duration of ice cover (days) 110 110 Mendota. and perspective to the dis- we very commonly ask in ecology. (C) there is a cycle of warmer winters recurring every few years (now known to be the 70 70 result of the El Niño Southern Oscillation. These opinions may be 90 passionately held and forcefully argued. As the evidence supporting a theory accu- 50 mulates. nonscientists will Second. while others will wait until 30 the bulk of the evidence is greater. see Chapter 18). During the process of amassing evi- Duration of ice cover (days) dence regarding the validity of a theory.3 130 130 Repeated observations over space or time can reveal infor- mation that is not apparent from one or a few observations.) 30 30 1998 1988 1998 processes of competition and herbivory each Warmer contribute to shaping this community?” So. We can ask whether a particular piece belongs in this spot. saying that it does not belong to any puzzle. We may even conclude that this particu- lar piece does not belong in this puzzle. also contribute to the debate and may be able to offer work fails to account for a second type of question that valuable insight. When we of scientists knowledgeable in that field are convinced examine the structure of a plant community. we ask about the theory becomes overwhelming and the vast majority relative importance of different processes. have been kept for more than 142 years. winters in Wisconsin are warmer now than 50 50 they were 142 years ago. Less 30 1950 1960 1970 1980 1990 2000 often are we attempting to completely Warmer throw away the piece. some scientists will be willing to accept it sooner. by erecting a hypothesis and falsifying 50 it. but a more limited one than Popper envis- 90 aged. Wisconsin. Theory construction is like assembling a jigsaw puzzle from a pile of pieces from 70 more than one box. “Is it true or false that competition is occurring?” startling new evidence or a new. will lead different scientists to differing opinions. and in what ways. The information for a single year (A) is fairly mean- ingless. when we are building our theories about Colder 50 Years 130 plant community structure. and different weight given to different pieces of evidence. Often the issue is not cussion. Duration of ice cover (days) 110 Falsification does play a role in science. As an example. as it does in all scientific fields. (After Magnuson et al. and (D) overall. 1860 1880 1900 1920 1940 1960 1980 2000 Warmer If the issue under debate has political or economic implications. and discussion can sometimes become heated. do the a reevaluation). Colder 142 Years Controversy also plays an important 170 part in ecology. our activities are more akin to assembling a complex model than to falsifying a set of propositions. 2001. PAGE PROOF: 2ND PASS The Science of Plant Ecology 7 Colder 1 Year 10 Years Figure 1. Rather. broader theory forces Instead. and more importantly. it is the judgment of scien- .

Y has a 90% chance of extinction within the next 20 years—that does not necessarily dictate any particular Scale and Heterogeneity public policy. and expanding one’s focus to more of a particular study to other organisms or places. for example. giving used to evaluate where the outcome of a particular them equal weight would be contrary to the way science experiment fits in with—or differs from—the results of works. and on what costs A great deal of recent interest in ecology has been gen- they are willing to pay to do so. This approach has issues of public policy. and so on they would tell us nothing beyond something of such up to global-scale phenomena. or to other pairs of species nental scale. there would be no at the level of organisms to those that function at a scale value and no point in doing any experiments. that if more evidence for a number of important ecological questions than x% of its remaining habitat is lost. watersheds. determined by patterns of glaciation ments involving helium deal with a universal entity. helium atom. This is the pop. local community.g. it is unreasonable to consider that theory workings of existing models. For to entire organisms. known collectively as meta-analysis. ular species at a distance from one another. Likewise. These supported by evidence are not the same as those sup. The same phe- species Y could use the scientific conclusions for their nomenon can be seen very differently when studied own ends. A different kind of hierarchy could this reason.8 Chapter 1 PAGE PROOF: 2ND PASS tists that must decide the answers to scientific questions. 2001). within a small local area and at the scale of a landscape clusions to promote conservation. we might notice that the plants are grouped quarks. processes vary as a function of the scale at which they one who wanted (for whatever reason) to exterminate operate and are studied (Figure 1. Policy decisions depend on how impor- tant people think it is to save species Y. (see Figure 1. and one can move up and down many dif- (e. . Opinions not sis of the results of independent experiments.. In the than one scale can be richly rewarding. At a larger tons and two neutrons. At a regional or conti- to other seasons or locations. While we personally erated by consideration of how ecological patterns and hope this would never happen. arated enclaves. tations of plants over a range of different environments tremes. nated. which in turn are made up of scale. For instance. together across the landscape because individuals that ular image of scientific theories. communities to ecosystems and up to entire biomes. That may work for approach is to use methods for the quantitative synthe- the democratic process. One approach to resolving this tension is to see how When a scientific consensus has been reached on a sci. In a study of a fields of chemistry or physics. just as we could use those same scientific con. We often refer to these scale changes in terms of a petition. Another everyone’s opinion is equally valid. extremely cautious scientists take the posi. can be ported by the weight of a great deal of evidence. with no qualifications needed. ogy. we recognize that some. Different kinds of things may be going tion arises as to how far one can extend the conclusions on at different scales.. in experiments on plant com. for instance. essarily congruent: one might study the individual adap- The truth is somewhere between these two ex. or region—that is. In contrast. then plant species in recent years (Gurevitch et al. If that were so.g. expand from individual organisms to populations to tion that no conclusion can be extended beyond the par. These levels are not nec- limited scope as to be worthless. at different spatial scales. and the surroundings change as well can move from the level of molecules to tissues to organs (e. methods. regions. or because the seeds iment on competition between two plant species extend have limited ability to disperse. because of habitats. ferent kinds of hierarchies in ecology. For example. Do the results of a field exper. we might see that com- ment are considered to be absolutely true for all times petitive interactions keep individual plants of a partic- and places: an atom of helium is made up of two pro.3). landscapes. across an entire landscape or even a region. if scientists are in been used to evaluate the broad body of experimental strong agreement about something—say. the weather is different this year than last year). the ques. an ticular conditions that existed when the experiment was alternative hierarchy might move from things that occur conducted.4). one ically identical). one’s perspective can change dramatically when study- Specific Results versus General Understanding ing an ecological process over a single growing season Because ecologists work at such a variety of scales and of a few months or over a period of decades or centuries on such a diversity of organisms and systems. however. but not in science. the individual plants used each time are not genet. are too far apart from any others never become polli- Ecology is different. other similar experiments conducted on different organ- This does not mean that scientists should decide isms at different places and times. and fail to leave descendents. creating a constant and dynamic tension in ecol. and whether it supports or to be just another guess or opinion and to hold that rejects the predictions of those models. the plants may exist in several large but sep- within the same families or functional groups? Experi. the exact composition of the entities changes hierarchy. the outcome of a particular experiment fits into the entific theory. the and species migration thousands of years in the past. the results of an experi.

per- the average conditions in the general habitat (Figure 1. and at a longer scale as climate changes over thou- sands of years all have important influences on plants. But the con. climatology. such as the diffusion of CO2 molecules in a leaf. or it may be the interplay between process- One of the reasons that scale is now recognized as es occurring at different scales (such as CO2 flux in a for- being so important is that the world is a very heteroge. ecologists 10–2 B study patterns that stretch across the entire globe (105 m) C and thousands or even millions of years (103–106 or more A years). PAGE PROOF: 2ND PASS The Science of Plant Ecology 9 106 Figure 1. behavior. est canopy over the course of a day or a season). Finally. mental influences over larger areas and over generations lar concern in plant ecology because plants cannot move.4 H Ecologists study patterns and processes across a wide range I of scales in space and time. sometimes average these sorts of microenviron- ing organisms. Like. to changing conditions by evolving or by changing their duction. The habitat of a population or species is the kind of environment it generally inhabits and includes the set The Structure and History of Plant Ecology of biotic (living) and abiotic (nonliving) factors that influence it in the places one usually finds it. Varia- tions in conditions from day to night. and other organisms branches of mathematics. size. the with include geology. geography. as fluctuations in light levels in a small forest gap on a wise. or it may be large-scale. the environment critical. genetics. pollinators. evolutionary biology. we enter the realm of populations and communities and their changes D over the course of years and decades. mature terrestrial plants generally are firm. or how population interactions at very local scales con. occur over the shortest distances (10–4 m) and times (10–7 years). haps to a greater extent than does any other subject. By that we do not ditions in the immediate surroundings of an individual mean that it is unnatural or artificial. long-term time period may be completely upended when the same average conditions (such as CO2 concentration in the data are examined for trends over longer periods of time. the environment varies from moment to A CO2 transport in leaves F Population moment. Ecology is a very synthetic subject. summer to win- D Annual plant (growing phase) I Climate change. reproduction. At even larger scales. neous place. There are no specific ecological terms for the B Nutrient transport in soil G Community components of temporal heterogeneity. Even over very small distances. heterogeneity again becomes dispersed some distance away. growth. such as populations and can change in ways that may be very important to liv. the domain of the whole plant and 102 its birth.5). As continents are carried apart on tectonic plates immediately surrounding an individual plant is over. G F Moving up the hierarchy. ter. So. one’s interpretation of data collected over a short partially cloudy day). distributions. at least. although their offspring may be time. developmental biology. Depending on the ecological process being studied and the organisms involved. or else become extinct. and mutualistic or pathogen- Time (years) ic fungi or bacteria). systematics. atmosphere). the presence. conditions Groups of organisms. paleoecology. 10–4 10–6 10–4 10–2 100 102 104 106 (1 year) seed eaters or dispersers. effects of heterogeneity. ology. At yet 100 E larger scales (101–102 m and 101–102 years). and death encompasses Space (m) slightly longer distances (100 m) and times (100 years). and climates are altered. of organisms’ lives. Similarly. species. or snowy E Perennial plant (growing phase) continental drift years. statistics and other and identity of neighboring plants. range shifts at many scales and has major effects on plants. growth and repro. ence. physi- in the immediate surroundings (grazers. and . but that it brings plant—its microhabitat—may differ considerably from together a very wide range of other fields of science. Factors operating to distinguish a microhabitat from oth. moment-to-moment variation that matters most (such tribute to the global range limitations of a species. Environmental conditions are a particu. This averaging acts to counter the Or. it may be the small-scale. The processes of plant physiolo- 104 gy. cold years. Some of the fields that ecology encompasses or overlaps ers around it include the composition of the soil. particularly over evolutionary ly rooted in place. molecular biology. soil sci- microclimate of the immediate area. across periods of wet years. organisms must either respond whelmingly important to its survival. but it also exists C Seeds H Biomes.

Individual plants experience the conditions in their immediate microhabitat. and by early in the English-speaking scientific community. carried out by ecology and some of the key figures in that history is professional and amateur naturalists in Europe and given in Chapters 12 and 13. 1993). and the story of his five-year voyage was pub- organisms in nature. Charles Darwin was one such ship’s nat- the ways in which ecologists think about and study uralist. when people’s health and survival depended on for these two subfields are synecology and autecology.5 The environment in a particular microhabitat can differ in a number of ways from con- ditions in the surrounding area. distinct traditions and histories. discuss particular topics and subfields. Instead. ogists and botanists focused on whole communities. Some early plant ecol- Westman and Peet 1985. warmer (more heat from sun) North-facing. biochemistry. Plant community ecologists. not the average conditions in the general area. some of which have quite and papers describe parts of its history (McIntosh 1985.6). North America and in their travels throughout the Early studies in plant autecology were especially world. several books ber of different subdisciplines.10 Chapter 1 PAGE PROOF: 2ND PASS N Sun 1m Exposed to wind South-facing. remarkable range of organisms and environments they it of an ecologist and how familiarity with them affects encountered. most notably Aristotle. were active in gy of plants with great accuracy. often carried a ship’s naturalist who cataloged the out this book. The modern science of during the first half of the twentieth century. we sketch some nineteenth century. their abilities to understand many aspects of the ecolo. with an admitted bias toward el coined the term “oecology” in 1866. than on the ridges. here. the twentieth century the Ecological Society of Ameri- torical details are scattered throughout the book as we ca had formed. a recognized discipline coalesced in the latter half of the itive history of plant ecology. in the the nineteenth century and dominated plant ecology fourth and fifth centuries B. while others focused on single species and the proper- The roots of plant ecology go back to prehistoric ties of individuals. lished as his Voyage of the Beagle (Figure 1. Other his. A more plant ecology began as the study of natural history in detailed discussion of the history of plant community the eighteenth and nineteenth centuries. Other potential effects of a microclimate are also illustrated. Ecology as a science the origins of ecology as a discipline in the last part of began with the Greeks. The older (now largely archaic) terms times. in particular. The German biologist Ernst Haeck- of the major milestones. Ships on journeys of discovery and colonization concerned with understanding unique plant adaptations . and per- haps greater herbivory due to its proximity to the groundhog burrow. The grass on the north-facing slope at the left of the diagram experiences cooler temperatures. showing you how they fit into the toolk. and competition for light and soil moisture may be more intense.C. Allen et al. Nicholson 1990. While no single Plant ecology as a discipline is made up of a num- definitive history of plant ecology exists. the soils are moister. The plants in the low-lying area may experience frost sooner in the fall and later in the spring than the surrounding areas due to cold air drainage.E. burrow more competition Subsurface drainage Moister soils Low-lying area Figure 1. Ecology as This is not the place to present a detailed and defin. We touch on many of these fields through. Cold air drainage Re-radiation Drier cooler of heat ridges Greater herbivory Water Slope Groundhog runoff Frost pockets Dense vegetation.

The second develop- quarters of the twentieth century developed independ. Two related developments were ogy or plant population ecology as far back as the nine. continue to this day. plant physi. Undoubtedly this was because fields of plant physiological and functional ecology. Animal community ecology has aspects of ecology in the 1970s and 1980s. responsible. This is et al.) to extreme environments. On the Origin of Species. for instance. further advances in technology made it possible for Conversely. Inc. and herbivory (see Chapter 11). Although some major insights were Plant population ecology drew on these ideas and the- gained. plant ecology for the first three. 1831 on a 5-year mission to chart the oceans and collect biological information from around the world. be shoehorned into many of the theories constructed for Beginning in the middle of the twentieth century. shortly after com- pleting the voyage. Substantial work in animal popula. plants and animals. which tran- a long history parallel to that of plant community ecol.6 H. Darwin collected vast numbers of plant and animal specimens and recorded copious scientific observations that were instrumental in the creation of his most famous work. . autecology began to fission into and their environments easier to characterize. although distinct subfields that today we would label as plant physiological ecol. scended the traditional separation of the studies of ogy (Mitman 1992). Lotka.S. animal subdiscipline had its origins in Great Britain in the physiological ecology joined with evolutionary biolo- 1960s. especially pollination (see Chapter 8) For the most part. and others). At the characteristics of plants are much easier to measure. physiological ecology advanced earlier physiological studies to come out of the greenhouse and and more rapidly among plant ecologists than it did into nature. about the same time. of animals (for most purposes. On the other hand. than those subfields that focused on single individuals and on pop. he is pictured here at the age of 24. events. nature of plants made it obvious they could no longer trolled laboratory environments. Beagle sailed from England December 27. one does not have to ulations. Plant population ecology as a recognizable catch plants!). Eventually new the- methodology became more sophisticated. mal interactions. in the 1980s. eventually leading to the creation of the among animal ecologists. (Images from Science Photo Library/Photo Researchers. ories were needed as discoveries about the unique ologists began to carry out most of their research in con. and a number tion ecology extends back to at least the 1920s (for exam- of famous studies were concerned with plant perform. a number of individuals in many The gap between the fields of plant and animal ecol- countries around the world were carrying out studies ogy was bridged in the 1970s. ple.M. 1987). PAGE PROOF: 2ND PASS The Science of Plant Ecology 11 Figure 1. ance in the field. animals. a move that plant biologists have not yet necessarily a very simplified and limited description of clearly made. ment was the burgeoning interest in the evolutionary ently of animal ecology. technological limitations severely hampered the ories as it was developing. as well as other ideas that development of the field. such as deserts. gy to create the field of evolutionary physiology (Feder It then spread to North America in the 1970s. Pearl. particularly with John Harper and his students. The first was the rise of studies of plant-ani- teenth century. with the work of Gause. As instrumentation and originated among plant ecologists. Charles Darwin sailed with the Beagle as ship’s naturalist.

. ogy. University of Chicago Press. These studies may cover a range from gy have seen major shifts in emphasis. Nat. D. 122: 697–705.12 Chapter 1 PAGE PROOF: 2ND PASS The most recent changes in the field of plant ecolo. Platt. Chica- ing. Chicago. Jones. G. Conservation of ecosystem ecology. Strong inference. community. J. R. Additional Readings Classic References Mayo. Ecological Understand- Hull. more integrated research vation ecology as recognized disciplines in the late 1980s. now. R. Previously it was dominated by questions about aries among subdisciplines. The Background of Ecology. systems. A. molecular genetics up through ecosystems and social ty ecology has seen such a shift in the past quarter cen. Rev. projects that involve many collaborators and examine Landscape ecologists began their careers from various phenomena across large scales of space and time or different directions. A major trend in contemporary ecology. Additional Resources McIntosh. Academic Press. Error and the Growth of Experimental Knowledge. including gy have been the rise of landscape ecology and conser. Am. J. K. The Logic of Scientific Discovery. Contemporary Research Pickett. excitement in this book. Kolasa and C. G. 1996. its main ing exciting times. M. 1985. recent years. about interactions within and among species. Plant ecology is experienc- whole-community patterns and processes. Hutchinson & Co. 1983. San Diego. IL. Cambridge University Press. London. 56:91–161. Salt. 1959. and are erasing many of the traditional bound- tury. Cambridge. W. R. 1988. L. 1964. investigator studies were very rare in ecology until and ecosystem ecology.. Bot. Henry Allan Gleason and the individualistic hypoth- Popper. Nicholson. including fields as diverse as plant across levels of organization. and we hope you will sense that focus has shifted to questions closer to population ecol. CA. plant ecology. such multi- in mathematical modeling and population. is toward larger. IL. go. 1994. Roles: Their limits and responsibilities in ecological and evolutionary research. G. T.. P. Science as a Process. D. Other fields within plant ecolo. esis: The structure of a botanist’s career. which was undertaking projects ecologists likewise created their field from backgrounds with large teams of scientists in the 1970s. Except in the subdiscipline community ecology and remote sensing. Uni- versity of Chicago Press. 1990. Science 146 : 347–353. S. Plant communi.