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842<br />

Chapter | 28 Avian <strong>Clinical</strong> <strong>Biochemistry</strong><br />

Plasma glucose (mmol/l)<br />

20<br />

18<br />

16<br />

14<br />

12<br />

0 0°° 8°° 12°° 16°° 20°° 24°° 4°°<br />

Time <strong>of</strong> day<br />

FIGURE 28-2 Mean ( SEM) plasma glucose concentration as a function<br />

<strong>of</strong> time in fasted racing pigeons. A cosine function y(t) is fitted to<br />

the data: y(t) 16.26 1.55 • cos (0.2618t to 2.4646). The relevance <strong>of</strong><br />

the fit as judged by means <strong>of</strong> the multiple correlation coefficient was significant<br />

(R 0.892, p 0.01). Reprinted with permission from Lumeij<br />

et al . (1987b).<br />

flights or to incubate eggs ( Blem, 2000 ). During migratory<br />

flights, energy expenditure may be more than seven<br />

times the basic metabolic rate for 3 to 5 days ( Battley,<br />

2003 ). Lipid reserves in birds are stored as triglycerides,<br />

which have a high caloric density and do not require water.<br />

Storage <strong>of</strong> fat reserves may double the body mass in some<br />

cases.<br />

Starved birds go through three phases. The total duration<br />

<strong>of</strong> the various phases depends on the size <strong>of</strong> the bird<br />

and the amount <strong>of</strong> lipid reserves and may vary from a few<br />

hours in hummingbirds to 5 months in emperor penguins<br />

( Robin et al ., 1998 ).<br />

In phase 1, food in the digestive tract and glycogen<br />

reserves are used; in phase 2, lipids are metabolized; and<br />

in phase 3, protein is used as a substrate for glucose synthesis.<br />

Phase 2 <strong>of</strong> starvation is characterized by an almost<br />

constant rate <strong>of</strong> body mass loss, a low respiratory quotient,<br />

and steady plasma concentrations <strong>of</strong> uric acid and β<br />

hydroxybutyrate. Although fatty acids provide the main<br />

energy source, about 5% <strong>of</strong> the energy is provided by protein<br />

breakdown, to generate citric acid intermediates, to<br />

act as substrate for gluconeogenesis, and for production<br />

<strong>of</strong> antioxidants ( Battley, 2003 ). When critical depletion <strong>of</strong><br />

fat stores is imminent, phase 3 <strong>of</strong> starvation is heralded by<br />

lowering <strong>of</strong> plasma concentrations <strong>of</strong> β-hydroxybutyrate<br />

and increased uric acid concentrations, reflecting a<br />

decreased contribution <strong>of</strong> lipids to energy metabolism and<br />

increased protein catabolism, respectively ( Fig. 28-3 ). The<br />

refeeding drive is related to the attainment to a given energy<br />

status rather than to a given duration <strong>of</strong> fasting or body<br />

mass loss ( Robin et al ., 1998 ). The metabolic shift to an<br />

increased protein breakdown is regulated by an endocrine<br />

level, nmol·L 1 level, mmol·L 1 display song<br />

Corticosterone Plasma b–OHB<br />

Behavioral<br />

index and<br />

Body mass<br />

Plasma uric<br />

loss<br />

g·kg 1·24h acid level,<br />

1<br />

3<br />

2<br />

1<br />

0<br />

1.0<br />

0.8<br />

0.6<br />

0.4<br />

90<br />

60<br />

30<br />

0<br />

1.8<br />

1.2<br />

0.6<br />

0.0<br />

24<br />

18<br />

12<br />

6<br />

0<br />

mmol·L 1 Plasma<br />

36 33 30 27 24 21 18<br />

Body mass, kg<br />

FIGURE 28-3 Changes in specific daily body mass, plasma uric acid,<br />

corticosterone, and β-hydroxybutyrate (β -OHB), and behavior versus<br />

body mass in spontaneously fasted emperor penguins (X SE; n 6).<br />

Crosshatched bars represent periods <strong>of</strong> display songs. The behavioral<br />

index was calculated as number <strong>of</strong> days an animal was active during<br />

successive periods <strong>of</strong> fasting corresponding to a 2-kg loss in body mass.<br />

Reprinted with permission from Robin et al . (1998) .<br />

shift (elevated corticosterone concentration), after which<br />

a further adrenocortical response to an acute stressor is<br />

inhibited. The adrenocortical response typical for an emergency<br />

situation is only reached when muscle protein is<br />

dangerously low ( Jenni et al. , 2000 ) .<br />

Mortality resulting from hepatic lipidosis has been<br />

described in a wide variety <strong>of</strong> avian species ( James et al .,<br />

2000 ; Wadsworth et al. , 1984 ) including, among others,<br />

chickens ( Butler, 1976 ), turkeys ( Gazdinski et al. , 1994 ),

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