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Vergara - 1976 - Physiological and morphological adaptability of ri

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348 crLnuATs AND RICE<br />

Expe<strong>ri</strong>mental studies on insect bioclimatics. on the other h<strong>and</strong>. are those in<br />

which the insect in question is exposed to controlled climatic va<strong>ri</strong>ables in the<br />

laboratory or in the field, <strong>and</strong> conclusions about the insects bioclimatic characte<strong>ri</strong>stics<br />

are drawn directly from the results <strong>of</strong> the expe<strong>ri</strong>ments. Constant teinpcrature<br />

incubators. environmental control chambers or bioclimatic chambers<br />

are the p<strong>ri</strong>ncipal pieces <strong>of</strong> equipment used for such bioclimatic studies. These<br />

are the entomological equivalents <strong>of</strong> phytotrons. or plant growth chambers.<br />

Climates are composed <strong>of</strong> a large number <strong>of</strong> factors involving aspects <strong>of</strong> atmosphe<strong>ri</strong>c<br />

heating <strong>and</strong> cooling (insolation, radiation); sunlight intensity". air<br />

movement (wind); moisture content <strong>of</strong> the air (relative or absolute humidity.<br />

saturation deficit); moisture condensation from the air (rain. snow. hail, ice.<br />

dew); cloud <strong>and</strong> overcast; baromet<strong>ri</strong>c pressure; percentage <strong>of</strong> sunshine. <strong>and</strong><br />

so on. In certain zones <strong>of</strong> the earth (temperate. subarctic, arctic). the length <strong>of</strong><br />

the growing season (frost-free days) is a measurable aspect <strong>of</strong> the seasonality <strong>of</strong><br />

climate, as are allemations <strong>of</strong> wet <strong>and</strong> dry seasons in a<strong>ri</strong>d, semia<strong>ri</strong>d, or monsoon<br />

regions.<br />

Not all <strong>of</strong> these individualized factors are important to any particular form <strong>of</strong><br />

life or occur at dominant intensities at a particular place. Those factors considered<br />

important in insect ecology are temperature, relative humidity, evapora~<br />

tion. precipitation in certain respects, wind. <strong>and</strong> photope<strong>ri</strong>od. Because <strong>of</strong> its<br />

very strong interaction with the insects physiological systems. temperature is<br />

probably the most important <strong>and</strong> widely studied climatic factor in insect bioclimatology.<br />

Because insects. particularly terrest<strong>ri</strong>al species. always have a<br />

problem <strong>of</strong> water balance. water retention, <strong>and</strong> evaporative water loss, relative<br />

humidity ranks close to temperature as an important limiting factor for insects.<br />

Photope<strong>ri</strong>od is a third, very influential factor. particularly for temperate or<br />

subarctic or arctic species, particularly those that must withst<strong>and</strong> winter cold or<br />

summer dryness. Thus. most expe<strong>ri</strong>mental studies <strong>of</strong> insects <strong>and</strong> climate involve<br />

one or more <strong>of</strong> these three major factors.<br />

The features <strong>of</strong> insect life that respond to or are most sensitive to climatic<br />

factors are numerous <strong>and</strong> complex. Biological systems that are influenced by<br />

climate are physiological, behavioral. <strong>and</strong> populational. With respect to temperature.<br />

practically every conceivable life process is affected, including development,<br />

hatching, melting, pupation. ecdysis, diapause, spennatogenesis, <strong>and</strong> ovigenesis.<br />

Behavioral processes affected include crawling. feeding, webbing.<br />

cocoon spinning, flight. courtship. mating, oviposition. <strong>and</strong> nest building. At the<br />

populational level, such aspects as birth rates. death rates, rates <strong>of</strong> increase in<br />

numbers. dispersal. <strong>and</strong> migration are all affected by climate. Any one <strong>of</strong> these<br />

features, physiological. behavioral. or populational. may be the single most<br />

important feature influenced by climate.<br />

For any one climatic factor. different factor intensities induce different<br />

amounts <strong>of</strong> response. At extremes <strong>of</strong> factor value, the typical response may be<br />

inhibited. This relationship has been called "Shelfords law <strong>of</strong> tolerance"<br />

(Odum. 1971). Using temperature. for example, there is a tolerance range <strong>of</strong>

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