Cm. Wood et al., PULSATILE UREA EXCRETION IN THE UREAGENIC TOADFISH OPSANUS-BETA - AN ANALYSIS OF RATES AND ROUTES, Journal of Experimental Biology, 198(8), 1995, pp. 1729-1741
This study focused on the rates and routes of urea-N and ammonia-N exc
retion in the ureagenic toadfish and on the possibility that urea-N ex
cretion occurs in pulses, Experimental approaches included the followi
ng: confinement in small individual containers with automated hourly s
ampling of water to follow temporal excretion patterns; divided chambe
rs to separate excretion from the anterior and posterior parts of the
fish; collection of urine and rectal fluid via chronic indwelling cath
eters; and gavage with [C-14]-labelled polyethylene glycol 4000 to det
ect regurgitation of gastrointestinal fluids, When a standardized 'cro
wding' pre-treatment was employed to induce ureotelic behaviour, the f
ish exhibited significant elevations in the activity of glutamine synt
hetase in liver, kidney and gills, elevated plasma and bile urea-N lev
els, but unchanged ammonia-N and urea-N levels in most other body flui
ds, Unencumbered ureotelic fish confined in small containers excreted
82% of their waste-N as urea-N and 18% as ammonia-N; almost all (94%)
of this urea-N excretion occurred in a single pulse of less than 3 h d
uration about once every 24h, This daily pulse did not occur by regurg
itation of gut fluids, by excretion through prominent pores behind the
pectoral fins or by discharge of rectal fluid or urine, Intestinal an
d urinary excretion accounted for less than 10% of whole-body urea-N e
xcretion and a negligible fraction of ammonia-N excretion, Pulsatile u
rea-N excretion occurred at the head end across the gills and/or body
surface, Ammonia-N excretion, which was not pulsatile, also occurred l
argely through the head end, However, once the toadfish had been place
d in divided chambers, urea-N excretion became continuous rather than
pulsatile, and ammonia-N excretion increased greatly, A severe stress
response was indicated by high levels of plasma cortisol, and the skin
, which lacks scales, became a significant route of both ammonia-N and
urea-N excretion, We speculate that the normal adaptive significance
is that ureotelism facilitates cryptic behaviour, allowing the toadfis
h to virtually eliminate N-waste excretion during long periods while i
t remains sheltered in burrows, However, during severe stress, the eff
ects of extremely high cortisol levels overwhelm the ammonia and urea
retention mechanisms, and both substances leak across the general body
surface.