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Encyclopedia of Evolution.pdf - Online Reading Center

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example <strong>of</strong> frequency-dependent selection is that <strong>of</strong> sex<br />

ratios (see sex, evolution <strong>of</strong>). If males are rare in a population,<br />

the average male will leave more <strong>of</strong>fspring than the<br />

average female. Conversely, rare females will leave more<br />

<strong>of</strong>fspring, on the average, than common males. Whichever<br />

sex is rare enjoys greater reproductive success. Frequencydependent<br />

selection is considered to be the main process by<br />

which most animal populations maintain an equal mix <strong>of</strong><br />

males and females.<br />

Limitations <strong>of</strong> Natural Selection<br />

Natural selection is not all-powerful. Below are examples <strong>of</strong><br />

limitations upon the power <strong>of</strong> natural selection:<br />

• Physical constraints. Natural selection cannot cause the<br />

evolution <strong>of</strong> characteristics that violate the laws <strong>of</strong> physics<br />

and chemistry. For example, natural selection cannot<br />

produce a large animal with skinny legs (see allometry).<br />

Also, the tallest trees in the world are about 350 feet<br />

(about 70 m) tall. Trees much taller than this are not possible,<br />

because there is a maximum height to which wood can<br />

conduct water, and the tallest redwoods may be very close<br />

to this maximum height. Large animals with wheels cannot<br />

evolve, for the wheel could not receive nutrition from the<br />

rest <strong>of</strong> the body through the axle.<br />

• Phyletic constraints. Once an evolutionary lineage has<br />

adopted a certain adaptation, this adaptation itself puts<br />

constraints on evolutionary change. It is not possible for<br />

an animal to have an exoskeleton and to be large. Once<br />

arthropods evolved exoskeletons (see invertebrates, evolution<br />

<strong>of</strong>), large size could no longer evolve in their lineage.<br />

• Absence <strong>of</strong> genetic variation. Natural selection can only<br />

choose from among the genetic characteristics that are<br />

available in the population. These characteristics arise by<br />

chance mutations.<br />

• Linkage <strong>of</strong> traits. Natural selection may be unable to select<br />

traits separately, if they are linked either on the same chromosome<br />

(as linkage groups; Mendelian genetics) or in<br />

development. In flowers, petal size and stamen size share<br />

developmental events, and natural selection cannot cause<br />

one to become bigger or smaller without the other.<br />

• Plasticity and dispersal. Organisms may respond to changes<br />

in environmental conditions by plasticity or acclimation,<br />

or by dispersal to a new location. These are ways in which<br />

individual organisms can avoid becoming the victims <strong>of</strong><br />

natural selection.<br />

• Post-reproductive individuals. Natural selection cannot<br />

eliminate harmful mutations that have their effect when all<br />

or most <strong>of</strong> the organism’s reproduction has been completed<br />

(see life history, evolution <strong>of</strong>). This is sometimes called<br />

the “kingpin principle,” based on a story about automobile<br />

manufacturer Henry Ford. He had his engineers study old<br />

Ford cars and find which part <strong>of</strong> the old cars was in the<br />

best condition. They reported that the kingpins were just<br />

like new. Ford told his factory managers to stop wasting<br />

their time making such good kingpins. There was no need<br />

for kingpins to last forever in a car that did not. Similarly,<br />

natural selection<br />

in organisms, there is no advantage for genes that promote<br />

the health and vigor <strong>of</strong> post-reproductive individuals.<br />

• The future. Natural selection cannot anticipate the future<br />

and prepare for it.<br />

• Group selection. Natural selection cannot favor characteristics<br />

that benefit the group but reduce the fitness <strong>of</strong> the<br />

individual that possesses them. Some adaptations result<br />

in harm for the individual (for example, worker bees that<br />

die after stinging a victim, and a mother spider that dies<br />

when her spiderlings eat her) but these are compensated<br />

by an increase in inclusive fitness (see altruism). When<br />

pathogens infect a host, natural selection may favor those<br />

selfish lineages <strong>of</strong> pathogens that reproduce most rapidly,<br />

even if the host dies as a result. Natural selection may<br />

work against such selfish pathogens if the death <strong>of</strong> the host<br />

results in the death <strong>of</strong> the entire pathogen population (see<br />

group selection) or if the illness <strong>of</strong> the host causes other<br />

hosts to stay away from the sick individual, thus preventing<br />

the dispersal <strong>of</strong> any pathogens <strong>of</strong> that population to new<br />

hosts (see evolutionary medicine). But natural selection<br />

cannot result in individuals sacrificing their own fitness<br />

within a population.<br />

Natural Selection Observed in Nature<br />

Below are just a few examples <strong>of</strong> natural selection that have<br />

been observed in nature.<br />

Stabilizing selection. <strong>Evolution</strong>ary biologists Arthur Weis<br />

and Warren Abrahamson studied insects that lay their eggs<br />

inside <strong>of</strong> plant stems. The grub releases a compound that<br />

causes the stem to produce a round swelling called a gall. The<br />

grub can develop, well fed, inside this gall. There are dangers<br />

from predators, however. Gall wasps can recognize galls and<br />

pierce their ovipositors into the gall, laying their eggs on the<br />

grub, and the wasp larvae eat the grub. Birds can also recognize<br />

galls and tear them open, to eat the protein-rich grub.<br />

Gall wasps prefer smaller galls; wasp parasitism therefore<br />

selects for larger galls. Birds prefer larger galls; bird predation<br />

therefore selects for smaller galls. The result is what looks<br />

like stabilizing selection for medium-sized galls but is actually<br />

a compromise <strong>of</strong> two processes <strong>of</strong> directional selection.<br />

Directional selection. Soapberry bugs (Jadera haematoloma)<br />

pierce fruits <strong>of</strong> certain plants with their long beaks.<br />

They are native to the keys <strong>of</strong> Florida, where they obtained<br />

their food from native balloon vines. After 1925, landscapers<br />

introduced the golden rain tree, an Asian relative <strong>of</strong> the balloon<br />

vine but with smaller fruits, into central Florida. Soapberry<br />

bugs began to use the new trees as a food source. In<br />

central Florida, natural selection favored bugs with shorter<br />

beaks. Over the next few decades, the central Florida bug<br />

populations evolved shorter beaks, while the bugs in the keys<br />

retained their long beaks. <strong>Evolution</strong>ary scientists Scott Carroll<br />

and Christin Boyd documented this change by examining<br />

bug collections that had been made before and after the<br />

introduction <strong>of</strong> golden rain trees to central Florida.<br />

Diversifying selection. In populations <strong>of</strong> African blackbellied<br />

seedcrackers (a type <strong>of</strong> bird), individuals have either<br />

large or small beaks. These two beak sizes correspond to two

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