Matters of Scale

Brian J. McGill
Science 328, 575 (2010);
DOI: 10.1126/science.1188528

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 PERSPECTIVES

in the eukaryotic genome. In comparison References e1000313 (2010).

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evolutionary processes. 9. International Aphid Genomics Consortium, PLoS Biol. 8, 10.1126/science.1190417

ECOLOGY
Recognition of the scale dependence of
Matters of Scale ecological processes helps explain the
distribution and abundance of organisms.
Brian J. McGill

Downloaded from www.sciencemag.org on June 18, 2012

I
n 1687, Newton reported that the same to solar systems. However, over the past 25 (6, 7). Now consideration of scale is helping
laws could describe Galileo’s data on years, ecologists have come to realize that, to address another key issue in ecology: the
balls rolling down ramps and Brahe’s data unlike physics, ecology is scale-dependent question of what controls the distribution and
on planets moving around the Sun (1). This (2–4). In a recent paper, Gotelli, Graves, and abundance of organisms. For example, why
observation implied that a finite list of princi- Rahbek (5) highlight the importance of this is the scissor-tailed flycatcher (Tyrannus for-
ples could explain our infinite universe. And scale dependence: They show that a process tificatus), one of North America’s most strik-
it inspired a leap across scales: The rules at that occurs at small spatial scales, namely ing birds, found mainly in Texas and Okla-
human scales are not unique. Newton’s laws competition between individuals, plays an homa? Four main factors limiting the distri-
of motion are still the dominant explanatory important role even at the large scale of an bution of species have been hypothesized.
tool across scales ranging from a few atoms entire country. Climate explains why the polar bear lives in
The realization that ecology is scale depen- the Arctic and palm trees grow in the tropics
School of Natural Resources, University of Arizona, Tucson, dent has recently helped to explain a multi- (8, 9). Random dispersal determines who can
AZ 85721, USA. E-mail: [email protected] tude of seemingly conflicting data in ecology get somewhere first or in large numbers. Spe-

What controls the distribu-

tion of species? Four main
processes (vertical axis) are
believed to control the dis-
tribution of organisms; their
relative importance changes
Climate with scale (horizontal axis).
The thickness of the bar for a
given factor at a given scale
indicates how important that
factor is at that scale. Ecolo-
gists began drawing such dia-
Species interactions grams 25 years ago (16), but
have only recently begun to
perform empirical studies to
test the suggested relation-
ships. The question mark at
intermediate scales of disper-
Habitat sal indicates that little data
exist on this process at these
? scales. Climate is impor-
tant for two scales, through
two processes: microclimate
PHOTO CREDITS: JUPITERIMAGES

(such as sun or shade) at

Dispersal small scales and biogeogra-
phy at large scales. Most ecol-
100 101 102 103 104 105 106 107
ogists will disagree with some
Most plot-based ecology
aspect of this figure, but it is
Denmark Biomes in North America the kind of complex, multi-
North America Earth (land) faceted, but testable hypoth-
Lineal scale (m)
esis that ecology needs.

www.sciencemag.org SCIENCE VOL 328 30 APRIL 2010 575

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PERSPECTIVES

cies interactions (competition, predation, and scales at which competition is important. dominate at large spatial scales usually occur
disease) determine whether a species thrives Gotelli et al. show that at the scale of a few over large temporal scales (2). Is this true?
or withers in a given environment (10–12). hundred kilometers on a side, competition And can the importance of different processes
The final factor is habitat: Cottonwoods grow is important, but we already know (13, 14) (the thickness of the bars in the scale diagram)
throughout the southwestern United States, that at the scale of a biome (roughly 1000 be measured quantitatively? Statistical tech-
but only along rivers. Which of these factors km by 500 km in the two cases studied), niques and nested sampling designs that tell
are most important? competition is not very important (see the us how much variation occurs in the variable
It is becoming clear that the answer figure). This is an astonishingly precise of interest at each scale could help to address
depends on scale. Competition is played out scale-dependent statement of when compe- these questions (15). The answers will help to
at small scales through interactions between tition is important and unimportant. put ecology on a more quantitative footing.
individual organisms (birds in this case). It Thus, Gotelli et al. provide an example of References
is difficult to imagine how the interaction how ecology can proceed. Rather than debat- 1. I. Newton, Philosophiæ Naturalis Principia Mathematica
between two birds can be influential at large ing which of the four forces is most important (S. Pepys, London, 1687).
2. S. A. Levin, Ecology 73, 1943 (1992).
scales, and indeed there is evidence that the in general, ecologists need to ask which force 3. J. A. Wiens, Funct. Ecol. 3, 385 (1989).
role of competition drops off to close to zero (or forces) is most important at a given scale 4. D. C. Schneider, Bioscience 51, 545 (2001).
at biome or nearly continental scales (13, 14). (see the figure).The first step toward identify- 5. N. J. Gotelli, G. Graves, C. Rahbek, Proc. Natl. Acad. Sci.
U.S.A. 107, 5030 (2010).
But there is a big gap between small (up to ing scale dependencies of this kind is to col-
6. M. L. Rosenzweig, Species Diversity in Space and Time

Downloaded from www.sciencemag.org on June 18, 2012

hundreds of meters) and large (thousands of lect more data on what controls species distri- (Cambridge Univ. Press, Cambridge, 1995).
kilometers) scales. Where exactly does com- bution and other variables (such as richness, 7. C. Rahbek, G. R. Graves, Proc. Natl. Acad. Sci. 98, 4534
petition disappear? productivity, and abundance) across scales. (2001).
8. G. Caughley, J. Short, G. C. Grigg, H. Nix, J. Anim. Ecol.
Gotelli et al. assembled an impressive However, this will lead to many distinct scale
56, 751 (1987).
data set on the distribution of birds at the diagrams such as that in the figure, one for 9. T. Root, J. Biogeogr. 15, 489 (1988).
scale of a country (Denmark). Based on the each variable to be explained. This raises sev- 10. R. H. MacArthur, Geographical Ecology: Patterns in the
evidence and thinking just mentioned, they eral new challenges and questions. Distribution of Species (Princeton Univ. Press, Princeton,
NJ, 1972).
expected that competition would no longer What is the minimum number of scale dia- 11. G. E. Hutchinson, Cold Spring Harb. Symp. Quant. Biol.
be influential at this scale, and that habitat grams that we need? Can we, for example, 22, 415 (1957).
(specifically, the varying types of vegeta- collapse the richness-area and richness-pro- 12. J. Silvertown, M. E. Dodd, D. J. G. Gowing, J. O. Mount-
ford, Nature 400, 61 (1999).
tion) would be most important in controlling ductivity diagrams into one? Given that scale 13. R. Russell, S. A. Wood, G. Allison, B. A. Menge, Am. Nat.
where bird species live. Surprisingly, they is relative to organisms—forces acting at a 167, E158 (2006).
found that habitat appeared unimportant, scale of 1 m are unlikely to be the same for 14. J. A. Veech, J. Biogeogr. 33, 2145 (2006).
but that competition was important in deter- bacteria and elephants—how can we rescale 15. B. J. McGill, Am. Nat. 172, 88 (2008).
16. A. Shmida, S. Ellner, Vegetatio 58, 29 (1984).
mining which bird species lived where. depending on the organism? Another factor is
The results help to put a band on the time. It has been suggested that processes that 10.1126/science.1188528

ASTRONOMY

Hidden Growth of Supermassive Changes in the fraction of quasars hidden

by gas and dust over cosmic time helps
confirm models of the evolution and
Black Holes in Galaxy Mergers merger rates of galaxies.

Joel Primack

B
lack holes are found at the centers of obscures our view of galactic centers in the axies is still streaming through the universe
massive galaxies. Although no light visible to x-ray regions. On page 600 of this and can be detected in some form. Some of
escapes from them, their presence can issue, Treister et al. (1) present an analysis this radiation is altered. For example, red-
be revealed by the glow of surrounding gases of data from several space-based telescopes, shifting occurs because the wavelengths of
compressed and heated by the driving force of showing that a greater fraction of quasars that photons stretch as the universe continues to
the black hole’s gravitation. This quasar emis- formed in the early universe were obscured expand, and some short-wavelength photons
sion ranges from low-energy radio waves to by dust, compared with its later stages. This like x-rays and ultraviolet light are absorbed
the highest-energy gamma-ray region of the is consistent with observational evidence on by dust and re-emitted at longer wavelengths.
electromagnetic spectrum. Quasar forma- the evolution over cosmic time of gas-rich To figure out what happened in the cosmic
tion can be driven by galaxy mergers, which galaxies and a theoretical model for the rate past, we must see the entire electromagnetic
change the distribution of gas around the at which they merge. spectrum, from the high-energy gamma rays
black hole. This process can also create stars Like geologists and evolutionary biolo- to the long-wavelength radio waves. Fortu-
that supernova and create interstellar dust that gists, astronomers reconstruct the past to nately, NASA’s Great Observatories in space
understand the present. Landforms erode and cover much of this wavelength range—x-rays
Physics Department, University of California Santa Cruz, only a tiny fraction of organisms fossilize, but (the Chandra X-ray Observatory), near ultra-
Santa Cruz, CA 95064, USA. E-mail: [email protected] all of the energy that was ever radiated by gal- violet to the near infrared (the refurbished

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