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Cockroache; Ecology, behavior & history - W.J. Bell

Cockroache; Ecology, behavior & history - W.J. Bell

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The most sophisticated pattern of nitrogen excretion<br />

occurs in at least nine species in the Blattellinae (Parcoblatta,<br />

Symploce, Paratemnopteryx), which void discrete,<br />

formed pellets high in urate content. These pellets<br />

are distinct from fecal waste (Fig. 5.8), suggesting that the<br />

packaging does not occur by chance. The cockroaches<br />

store urates internally as well (Cochran, 1979a). The level<br />

of dietary nitrogen in relation to metabolic demand for<br />

nitrogen is the controlling factor in whether uric acid is<br />

voided (Cochran, 1981; Cochran and Mullins, 1982;<br />

Lembke and Cochran, 1990). This is nicely illustrated in<br />

Fig. 5.9, which shows urate pellet excretion in female Parcoblatta<br />

fulvescens on different diets over the course of a<br />

reproductive cycle. Excreted urate pellets serve as a type<br />

of external nitrogen storage system, which may be accessed<br />

either by the excretor or by other members of the<br />

social group in these gregarious species. Reproducing females<br />

have been observed consuming the urate pellets,<br />

and they do so primarily when they are on a low-nitrogen,<br />

high-carbohydrate diet. A female carrying an egg<br />

case was even observed eating one, although they do not<br />

normally feed at this time. This system allows the cockroaches<br />

to deal very efficiently with foods that vary widely<br />

in nitrogen content. High nitrogen levels? The cockroaches<br />

store urates up to a certain level, and beyond that<br />

they excrete it in the form of pellets. Nitrogen limited?<br />

They mobilize and use their urate fat body reserves. Nitrogen<br />

depleted? They scavenge for high-nitrogen foods,<br />

including bird droppings and the urate pellets of conspecifics.<br />

Nitrogen unavailable? They slow or stop reproduction<br />

or development until it can be found (Cochran,<br />

1986b; Lembke and Cochran, 1990).<br />

Implications of the Bacteroid-Urate System<br />

The bacteroid-assisted ability of cockroaches to store,<br />

mobilize, and in some cases, transfer urates uniquely allows<br />

them to utilize nitrogen that is typically lost via excretion<br />

in the vast majority of insects (Cochran, 1985).<br />

These symbionts thus have a great deal of power in structuring<br />

the nutritional ecology and life <strong>history</strong> strategies<br />

of their hosts. Bacteroids damp out natural fluctuations<br />

in food availability, allowing cockroaches a degree of independence<br />

from the current food supply. An individual<br />

can engorge prodigiously at a single nitrogenous bonanza,<br />

like a bird dropping or a dead conspecific, then<br />

later, when these materials are required for reproduction,<br />

development, or maintenance, slowly mobilize the stored<br />

reserves from the fat body like a time-release vitamin. The<br />

legendary ability of cockroaches to withstand periods of<br />

starvation is at least in part based on this storage-mobilization<br />

physiology. The beauty of the system, however, is<br />

that stored urates are not only recycled internally by an<br />

individual, but, depending on the species, may be transferred<br />

to conspecifics, and used as currency in mating and<br />

parental investment strategies. Any individual in an ag-<br />

Fig. 5.8 “Salt and pepper” feces of Paratemnopteryx ( Shawella) couloniana; male, right; female<br />

and ootheca, left. The pile of feces to the left of the ootheca shows the variation in color of the<br />

pellets. Some of these have been separated into piles of the dark-colored fecal waste pellets (above<br />

the female) and the white, urate-filled pellets (arrow). Photo courtesy of Donald G. Cochran.<br />

MICROBES: THE UNSEEN INFLUENCE 85

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