1 3
Mar Biol (2014) 161:1179–1193
DOI 10.1007/s00227-014-2409-7
ORIGINAL PAPER
Acid–base regulation in the Dungeness crab
(Metacarcinus magister)
Stephanie Hans · Sandra Fehsenfeld ·
Jason R. Treberg · Dirk Weihrauch
Received: 5 July 2013 / Accepted: 12 February 2014 / Published online: 14 March 2014
© Springer-Verlag Berlin Heidelberg 2014
consumption, were reduced in crabs acclimated to elevated
pCO
2
, demonstrating that either (amino acid) oxidation is
reduced in response to this particular stress, or nitrogenous
wastes are excreted in an alternative form.
Introduction
In many marine organisms, a rise in seawater pCO
2
(envi-
ronmental hypercapnia) causes an increase in body fluid
pCO
2
in order to maintain an outwardly directed gradient
for the passive excretion of metabolic CO
2
(Fabry et al.
2008; Melzner et al. 2009). Hypercapnic environments
can therefore lead to a respiratory acidosis, where the
extracellular pH decreases if the H
+
produced from the
hydration of excess CO
2
is not buffered in the extracellu-
lar fluids (Melzner et al. 2009). The decline in pH could
prove detrimental since a stable internal pH is essential for
many physiological and biochemical processes (Somero
1986; Riggs 1988; Wheatly and Henry 1992; Fabry et al.
2008). To counteract the internal pH drop active metazo-
ans, such as teleost fish and brachyuran crabs, are equipped
with mechanisms to excrete acid equivalents and/or create
an influx of bases from the surrounding medium (Boron
2004). Green crabs Carcinus maenas and Dungeness crabs
Metacarcinus magister exposed to medium- and short-term
hypercapnia for example actively accumulate HCO
3
-
in the
hemolymph to compensate for the resulting acidosis and
successfully restore hemolymph pH (Pane and Barry 2007;
Appelhans et al. 2012; Fehsenfeld and Weihrauch 2013).
In aquatic crabs, acid–base regulation occurs mainly in
the gills and uses many of the transporters also involved in
osmoregulatory processes (Henry and Wheatly 1992). Na
+
/
K
+
-ATPase in the gills of the hyper-regulating C. maenas
generates the sodium gradient between the cytoplasm of
Abstract Homeostatic regulation allows organisms to
secure basic physiological processes in a varying envi-
ronment. To counteract fluctuations in ambient carbonate
system speciation due to elevated seawater pCO
2
(hyper-
capnia), many aquatic crustaceans excrete/accumulate
acid–base equivalents through their gills; however, not
much is known about the role of ammonia in this response.
The present study investigated the effects of hypercapnia on
acid–base and ammonia regulation in the Dungeness crab,
Metacarcinus magister on the whole animal and isolated
gill levels. Hemolymph pCO
2
and [HCO
3
-
] increased in M.
magister acclimated to elevated pCO
2
(330 Pa), while pH
remained stable. Additionally, hemolymph [Na
+
], [Ca
2+
],
and [SO
4
2-
] were significantly increased. When challenged
with varying pH during gill perfusion, the pH of the arti-
ficial hemolymph remained relatively unchanged. Over-
all, ammonia production and excretion, as well as oxygen
Stephanie Hans and Sandra Fehsenfeld have contributed equally
to this work.
Communicated by H.-O. Pörtner.
S. Hans · S. Fehsenfeld (*) · J. R. Treberg · D. Weihrauch
Department of Biological Sciences, University of Manitoba,
190 Dysart Road, Winnipeg, MB R3T2N2, Canada
e-mail: Sandra.Fehsenfeld@umanitoba.ca
S. Hans
e-mail: umhans2@cc.umanitoba.ca
J. R. Treberg
e-mail: Jason.Treberg@umanitoba.ca
D. Weihrauch
e-mail: Dirk.Weihrauch@ad.umanitoba.ca
J. R. Treberg
Department of Human Nutritional Sciences, University
of Manitoba, 190 Dysart Road, Winnipeg, MB R3T2N2, Canada