Fwd: Living Energies mini-series - Energy, Productivity & Biodiversity

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27 Jan 2004 10:54:06 -0000

Living Energies mini-series - Energy, Productivity & Biodiversity

press-release

 

The Institute of Science in Society

Science Society Sustainability

http://www.i-sis.org.uk

 

General Enquiries sam

Website/Mailing List press-release

ISIS Director m.w.ho

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Energy, Productivity & Biodiversity

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Generations of ecologists have puzzled over the causes of biodiversity and its

relationship with productivity. Dr. Mae-Wan Ho investigates.

 

A fully referenced version of this article are posted on ISIS members’ website.

Details here.

 

“Why are there so many kinds of animals?”

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This was the question asked by distinguished ecologist Evelyn Hutchinson [1] in

1959, the centenary of Darwin’s Origin of Species, a question that has remained

as enigmatic today as it was then [2].

 

 

There are about a million described species of animals, three-quarters of them

being insects, of which disproportionately large numbers belong to the order

Coleoptera, or beetles. In contrast to land animals, there are far fewer species

in the sea.

 

 

Hutchinson considered a number of possible explanations. Could food chains or

feeding relationships suffice? If one supposes an energy conversion efficiency

of 20% at every link of the chain, and each predator being twice as big as its

prey, the fifth animal link will have a population of one ten thousandth (10-4)

of the first, which is about as long as it would get. Food chains could hardly

generate a great deal of biodiversity.

 

 

Natural selection isn’t going to help; an overly efficient predator will simply

eat itself out of prey, thus breaking the link and making itself extinct in the

process. While lengthening the chain is difficult, shortening the chain is not,

the most dramatic example is the whale-bone whale, which can feed largely on

plankton.

 

 

What about the diversity of terrestrial plants which provide a variety of

different structures - bark, leaves, flowers and fruits - for different animals

to feed on. A major source of biodiversity of land animals was indeed introduced

by the evolution of almost 200 000 species of flowering plants, and the

three-quarters of a million species of insects are a product of that diversity.

But then, why are there so many different kinds of plants?

 

 

Part of the answer is that instead of linear food-chains, nature is replete with

food-webs. Most predators eat more than one species of prey, which reduces the

danger that it will eat its prey and itself extinct. So, at least part of the

answer to why there are so many kinds of animals and plants is that biodiverse

communities are better able to persist than less diverse communities. And that

was the origin of the idea that complex ecosystems are more stable, which has

been hotly debated to this day. While it may be intuitively obvious that the

more flexible the links in the foodweb, the less likely they will break;

mathematicians find it extraordinarily difficult to represent such flexibility,

and more so, to agree what constitutes stability, let alone complexity [3].

 

 

 

Energy available?

**************

 

Going back to biodiversity, ecologists have long noticed that while a hectare of

tropical rainforest contains an estimated 200 to 300 species of trees, the same

area of temperate forest contains only 20-30 species. One hypothesis is that

diversity is ultimately determined by the amount of energy available to an

ecosystem. Support for this idea came from measures of productivity and

biodiversity in different ecological communities. Productivity is the rate of

production of biomass by an ecosystem, and is in general determined by the rate

of energy supply.

 

 

High proportions of land and freshwater species on earth do occur in the

tropics, which receive the highest amount of the sun’s radiant energy. Average

species richness increases from high to low latitudes and this has been

documented for a wide spectrum of taxonomic groups, including protists

(single-celled organisms), trees, ants, woodpeckers and primates, and for data

across a range of spatial resolutions [4]. Species richness also appears to

increase with energy, measured as mean annual temperature, and

evapotranspiration.

 

 

But that doesn’t seem to be the whole story. Relationship between diversity and

productivity was found to vary at different spatial scales [2]. At large

geographical scale, such as across continents in the same latitude, diversity

generally increases with productivity. At smaller local scales (metres to

kilometers), several different patterns emerge.

 

 

Early studies found biodiversity peaking at intermediate levels of productivity

in a unimodal curve (a curve with a single hump). More recent reviews came up

with a variety of relationships, with diversity increasing, decreasing or

remaining unchanged as productivity increases. Although some of these patterns

suggest that energy is causally involved, other factors may also be important,

such as environmental heterogeneity: spatial or temporal variation in the

physical, chemical or biological features of the environment.

 

Complexity of the environment?

************************

 

In a simple lab experiment [5], the bacterium Pseudomonas fluorescens was used

to test the relationship between environmental heterogeneity and diversity. This

bacterium is known to rapidly differentiate into distinct ‘morphs’ in different

microhabitats in unmixed culture vessels. One major morph flourishes at the

interface between air and the liquid growth medium, another does best in the

center of the culture vessel and a third occupies the bottom of the vessel. The

researchers found that there are further variations within each major morph, so

that a total of ten types can be distinguished. Shaking the vessel eliminated

environmental heterogeneity and, with it, diversity. With a gradient of

productivity, a unimodal diversity curve was obtained. In other words, diversity

increased with energy available up to a point, and then decreased as available

energy increased further.

 

 

Ecosystems typically consist of plants and animal species of vastly different

sizes, from big mammals to birds, insects and microbes in the soil, which would

use resource that matches their size. Thus, the more finely the species can

divide up space and resources, the more species can coexist in the same habitat.

But how best to represent this environmental heterogeneity?

 

 

Mark Ritchie from the University of Utah, Logan, in the United States, and Han

Olff in Wageningen Agricultural University, in the Netherlands, reasoned that

the distributions of habitat, food and resources often appear to be

statistically self-similar over three to four orders of magnitude. If so, their

volume or area can be described with fractal geometry [6].

 

 

A fractal is a structure that has dimensions in between the usual 1, 2 or 3; and

‘self-similar’ refers to the property that the structure appears the same over

many scales. Typical examples are fern leaves, branching blood vessels and the

coastline.

 

 

In a fractal environment, body size determines the abundance of food and

resources that a species perceives, and it sets limits to the similarity in body

size between any two species. Ritchie and Olff derived a body size ratio between

species of adjacent sizes that declines with increasing organism size. That in

turn predicts how diverse the community can be.

 

 

Thus, energy, productivity and environmental heterogeneity all appear to play a

role in creating biodiversity.

In the next article (“Why are organisms so complex?” this series), I shall show

how biodiversity and productivity are intimately linked through energy capture

and storage in a sustainable system.

 

 

 

 

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General Enquiries sam

Website/Mailing List press-release

ISIS Director m.w.ho

 

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