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Ecology: Context for Evolutionary Change
Energy
The Currency of Nature
 

Ecology is essentially the study of the economy of nature. The currency of this economy is energy. Energy allows organisms to run their metabolism, to grow, to produce offspring and to obtain more energy. Ecologists study the ways that species “make a living”: how they extract resources, maximize profits, minimize costs and compete with others. It is important to remember that species do not exist independently of other species. Ecosystems are made up of interacting populations of organisms and their physical environment; populations are linked by energy flow.


Trophic levels - feeding relationships and flows of energy
Ultimately, all of the energy used by all organisms comes from the sun[*]. Some organisms (such as plants, algae) use energy from sunlight to convert carbon dioxide and water into carbohydrates; that is, they make their food[#](Taiz & Zeiger 2002). For this reason, they are referred to as producersOrganism that synthesizes its own food, rather than consuming the tissues of other organisms.

Other organisms use the energy stored in the tissues of producers. HerbivoresOrganism that eats plants or algae, getting its energy from the 1st trophic level. (also known as primary consumers) consume plants or parts of plants. Some organisms feed on and break down particles of organic matter, such as the remains or wastes of organisms. Predators get their energy by killing and eating other animals. And parasites can infest any of these, extracting energy while (usually) keeping the host alive.

The specifics of these feeding relationships can be described as a food web[#](Molles 2004). More generally, organisms in a food web can be grouped into trophic (feeding) levels based on where they get their food (see Figure 1).

Producers are responsible for all of the energy that enters a food web, so they are grouped as the first trophic level. Trophic levels 2 and higher contain consumersOrganism which gets its energy from the tissues of other organisms., those organisms that feed on others.

If producers can support enough herbivores, another trophic level may be sustained - one made up of predators, which get their energy by eating animals from trophic level 2 or higher. Some ecosystems are productive enough (that is, they have enough energy in them) to support a fourth trophic level of top predators that eat from levels 2 and 3.

Two other groups take advantage of the energy sequestered by plants: decomposersOrganism that eats dead or decaying organisms and parasites.

The energy available in an ecosystem is limited to that which is photosynthesized by producers (often ~1% of solar input of energy)[#](Molles 2004). In fact, it is limited more than you might expect. About 96% of the biomass of producers becomes detritus, unavailable to consumers. What’s more, as energy moves among trophic groups, about 90% of the energy present in one trophic level never makes it to the next trophic level. Energy is used to fuel metabolism, respiration and heat production. Energy is stored or used for growth, and some is lost to parasites. These losses of energy limit the number of trophic levels in an ecosystem[#](Molles 2004).

Why is this important?
Remember that, within any population, more offspring are born than can possibly survive. Some die before or shortly after birth, hatching, or germination. Those that survive have another problem: They need energy. Energy is an absolute necessity. All life depends on it. At a minimum, energy is needed simply to survive and grow. Energy used for these purposes can be called 'operating costs'.

For example, an acorn (which contains a small ‘packet’ of energy from the parent tree to fuel germination) sprouts and grows roots, stems, and leaves. As it grows, the tree needs energy to produce more wood, roots, stems, and leaves each year. Photosynthesis uses energy to create and transport sugars, and roots need energy to take in and transport nutrients and water[#](Taiz & Zeiger 2002).

Any energy intake beyond that needed to cover operating costs can be stored or used for reproduction. We can call this 'profit'. Typically, organisms have enough energy ‘profits’ for both reproduction and storage. Short-lived species, such as annual plants, have no need to store energy; all profits can be used for reproduction.

Maximizing Profits
Any traits that reduce operating costs will also increase profits. Reducing wasteful spending and increasing efficiency are two ways to reduce operating costs.

For example, many animals that spend their lives in total darkness, such as in caves or underground, do not have functioning eyes[**]. Eyes require energy to build and maintain. If an individual has a mutation that disrupts the construction of eyes and if eyes are of no use, the energy that would have been used to maintain eyes can be used to increase reproductive success[#](Culver, et al. 1995).[***] Over time, the increased reproductive success of eyeless-types would increase their proportion in the population. Eventually, there would be no individuals with eyes, because eyes are a waste of energy.

A mutation that improves metabolic efficiency would similarly become abundant if the extra profits increase reproductive success for those with the mutation. A good example of increased efficiency comes from plants in hot, dry climates. Plant leaves have small openings (stomatesSmall openings (primarily) on the underside of leaves. Their functionis to allow CO2 into the leaf and the waste products of photosynthesis out,and to regulate passage of water vapor out of the leaf.) through which carbon dioxide enters. Photosynthesis uses energy from sunlight to convert carbon dioxide to carbohydrates, in a process called the Calvin Cycle. Photosynthetic reactions take place inside structures called chloroplastsA chlorophyll-containing structure found in algae and plant cells.Photosynthesis takes place here. (in tissues called mesophyll; see Figure 2), and in most plants, carbon dioxide enters the Calvin Cycle directly.

When stomates are open, carbon dioxide can enter and oxygen can exit. But there's a tradeoff: water can also escape. In hot and dry conditions, plants close their stomates to prevent water loss. As a result, levels of oxygen (produced by photosynthetic reactions) increase.

The Calvin Cycle runs most efficiently using carbon dioxide, but can also use oxygen. In fact, a key enzyme is more likely to bind to oxygen. With stomates closed, the Calvin Cycle will run out of carbon dioxide, and begin to use oxygen. When this happens, the cell uses more energy in making sugars, often causing a net loss of energy[#](Taiz & Zeiger 2002).

A relatively recent (in evolutionary time) adaptation has arisen in some plants - particularly grasses - from warmer regions, especially areas with low atmospheric carbon dioxide concentrations. In these plants, carbon is sequestered in mesophyll and moved into adjacent cells (called bundle-sheath cells). Segregated from the oxygen in mesophyll, the Calvin Cycle proceeds more efficiently because it uses only carbon dioxide[#](Taiz & Zeiger 2002). This newer type of carbon fixation/photosynthesis (known as C4) has apparently evolved independently numerous times from species with the more primitive C3 carbon fixation[#](Osborne & Beerling 2006).

Individuals can also possess traits that improve their ability to sequester energy. Examples among animals include stronger claws for digging or improved teeth for grinding or cutting. Changes in the digestive system also can save energy, either by reducing the energy needed to digest food or by increasing the amount of energy that can be extracted from each item of food. For example, mammals have trouble digesting plants because mammals lack the enzymes needed to break down cellulose, a sugar that makes up plant cell walls. buffalo and deer have a modified stomach in which plant material is processed multiple times (see Figure 3). This process increases both energy-use efficiency and energy intake[#](Vaughan, et al. 2000).

Profits can be saved, for example, for use during times when food is scarce or during periods of hibernation. The oak tree stores energy over winter to begin growing leaves the following spring (until they can begin photosynthesis).

Profits are spent on many aspects of reproduction. Plants spend energy on flowers, nectar, pollen and ovules, and then produce fruits and seeds.  Animals spend energy on sperm, or eggs and embryos. They usually have to find mates, and perhaps compete with others in order to mate.  Many species feed and protect their offspring. 

Individuals with higher profits have an advantage over competitors. Some species use the higher profits to produce more offspring.  Others devote more energy to fuel each offspring's growth. 

If genetic changes make more energy available for reproduction and thereby improve reproductive success, those genetic changes will become increasingly common.  The population evolves and adapts to its environment.

 

Beyer, HL, EH Merrill, N Varley & MS Boyce. 2007. Willow on yellowstone's northern range: Evidence for a trophic cascade? Ecological Applications 17:1563–1571

Root, T. 1988. Energy constraints on avian distribution and abundances. Ecology 69:330-339

Currie, DJ. 1991. Energy and large-scale patterns of animal- and plant-species richness. The American Naturalist 137:27-49

 
* Okay, not all. But very nearly all. Microorganisms in dark, oxygen-deprived regions of the oceans use hydrogen gas, hydrogen sulfide or methane, rather than sunlight, as a source of energy. This is known as chemosynthesis.
**It is significant that in nearly all cases these organisms retain some non-functioning vestiges of eyes. This is expected by (and is best explained by) evolutionary theory

***But see Jefferey, et al (2000) for an interesting and different mechanism that created eyeless types in the cavefish Astyanax.

 
 

Figure 1. Trophic levels and flows of energy

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Figure 2. Leaf Anatomy.

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Figure 3. The ruminant stomach

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Copyright © 2007 Michael Kreuzer, Jr.