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