THERMOREGULATION AT LOW TEMPERATURES

_Body Temperature_

Body temperatures in Figure 6 are those recorded during metabolic

measurements from animals equipped with surgically implanted,

temperature-sensitive radio transmitters. Each point was recorded

during the lowest level of oxygen consumption at each T_{a}. In both

summer and winter, T_{b}'s were lowest during metabolic measurements

at T_{a}'s around T_{lc}. At T_{a}'s below T_{lc}, T_{b}'s increased

(Figure 6), which is an unusual response. Under similar conditions,

other procyonids either maintain a nearly constant T_{b} or allow it

to fall slightly (Mueller and Kulzer, 1977; Chevillard-Hugot et al.,

1980; Mueller and Rost, 1983; Chevalier, 1985). For our raccoons,

confinement in the metabolism chamber at low temperatures must have

stimulated a greater than necessary increase in metabolic rate such

that heat production exceeded heat loss, which caused T_{b} to become

elevated.

[Illustration: FIGURE 6.--Relationship between body temperature and

chamber air temperature in summer (panel A), and winter (panel B):

captive females, open circles and solid lines; captive males, solid

circles and dashed lines. Solid vertical lines represent lower

critical temperatures.]

TABLE 5.--Regression equations describing oxygen consumption

(mL O_{2}.g^{-1}.h^{-1}) of _Procyon lotor_ at temperatures below

their lower critical temperature (I = x-intercept ( deg.C), n = number

of observations, R squared = coefficient of determination, SEE = standard

error of estimate for the y-intercept (a) and slope (b), X = chamber

temperature ( deg.C), and Y = oxygen consumption).

----------------+------------------------------------------------------

Season | SEE

and sex | -----------

| Equation (n) R squared a b I

----------------+------------------------------------------------------

Summer |

Trapped male | Y = 1.09 - 0.0281.X (30) 0.64 0.0353 0.0040 38.8

Captive male | Y = 0.97 - 0.0258.X (12) 0.91 0.0235 0.0025 37.6

Captive female| Y = 1.04 - 0.0251.X (29) 0.78 0.0288 0.0026 41.1

Winter |

Captive, |

both sexes | Y = 0.68 - 0.0193.X (36) 0.68 0.0157 0.0023 35.2

----------------+------------------------------------------------------

_Summer_

During summer, T_{lc} for male raccoons was 20 deg.C, whereas for females

it was 25 deg.C (Figure 2). Regression equations calculated to describe

oxygen consumption at T_{a}'s below T_{lc} are presented in Table 5.

For three groups of summer animals, slopes of regressions are

identical. This indicates that minimum conductances of these three

groups were equivalent. Intercepts of these equations are different,

which suggests a difference in metabolic cost of thermoregulation

between these groups (Figure 2); captive males had a lower intercept

than either trapped males (p<0.005) or captive females (p<0.05), but

there was no difference in intercepts of captive females and trapped

males. These regression equations, therefore, also were derived using

values of oxygen consumption expressed in terms of metabolic body mass

(Mellen, 1963). Relationships between intercepts of these equations

are different than those for regressions in Table 5. Intercept for

females was intermediate to, and not different from, those of the two

groups of males. However, captive males still had a lower intercept

than trapped males (p<0.025). Thus, in summer, thermoregulatory

metabolism was less expensive for captive than for trapped males, and

in spite of a 5 deg.C difference in their T_{lc}'s (Figure 2), captive

males and females had similar thermoregulatory costs.

Regression lines for three groups of animals in summer extrapolate to

zero metabolism at values equivalent to, or greater than, normal

T_{b}; 38.8 deg.C for trapped males, 37.6 deg.C for captive males, and 41.1 deg.C

for captive females (Table 5). Thus, all three groups had minimized

thermal conductance at T_{a}'s below T_{lc} (Scholander et al., 1950b;

McNab, 1980b). Minimum wet thermal conductance calculated for raccoons

in summer with Eq. 4 (Table 3) is numerically similar to these "slope"

values (Table 5), and it was, therefore, considered to be the best

estimate of C_{mw} for _Procyon lotor_ during that season

(0.0256 mL O_{2}.g^{-1}.h^{-1}. deg.C^{-1}).

_Winter_

During winter T_{lc} for both sexes decreased to 11 deg.C (Figure 3).

Regression equations of thermoregulatory metabolism for males and

females in winter are not different from each other in either slope or

intercept. These data, therefore, were combined into a single equation

(Table 5). Slope and intercept of this equation are both lower

(p<0.005 and p<0.05, respectively) than those for summer animals

(Table 5). Identical results were obtained from comparisons using

regressions derived from oxygen consumption expressed in terms of

metabolic body mass (Mellen, 1963). Thermoregulatory costs at any

temperature below 20 deg.C were lower for winter than summer animals

(Figures 2, 3).

TABLE 6.--Regression equations describing oxygen consumption

(mL O_{2}.g^{-1}.h^{-1}) of _Procyon lotor_ at temperatures below

their lower critical temperature in winter (A = females with radio

transmitters, B = females without radio transmitters, C = males,

I = x-intercept ( deg.C), n = number of observations, R squared = coefficient

of determination, X = chamber temperature ( deg.C), and Y = oxygen

consumption).

-----+--------------------------------------

Group| Equation (n) R squared I

-----+--------------------------------------

A | Y = 0.63 - 0.0158.X (10) 0.66 40.1

B | Y = 0.72 - 0.0226.X (11) 0.71 32.1

C | Y = 0.69 - 0.0200.X (15) 0.79 34.7

-----+--------------------------------------

[Illustration: FIGURE 7.--Relationship between body temperature and

time of day at various months of the year: captive females, open

circles; captive males, closed circles. Vertical cross-hatched

areas represent civil twilight.]

The regression line for _Procyon lotor_ in winter (Table 5)

extrapolates to zero metabolism at 35.2 deg.C, which is below normal T_{b}

(Figures 6, 7). This suggests that not all raccoons measured in winter

minimized thermoregulatory metabolism or conductances at T_{a}'s below

T_{lc} (Scholander et al., 1950b; McNab, 1980b). To assess this

possibility, data for these animals were divided into three groups:

(A) females with radio transmitters, (B) females without radio

transmitters, and (C) males (Table 6). Regression equations of

metabolism below T_{lc} were derived for each group, and based on

extrapolated T_{b}'s at zero metabolism, only the two females with

implanted radio transmitters (group A) minimized thermoregulatory

metabolism and conductance. Had animals in groups B and C also

minimized their thermal conductances, while retaining their measured

metabolic rates, their rates of heat production would have been

disproportionately higher than their rates of heat loss. Equation 4

predicts that under these conditions their body temperatures would

have been elevated to 42.0 deg.C and 40.4 deg.C, respectively. Thus, in order

to avoid such a large increase in body temperature, animals in groups

B and C increased their thermal conductances in preference to lowering

their metabolic rates. The regression equation of thermoregulatory

metabolism for all winter animals (Table 5), therefore, overestimates

minimum metabolic cost of temperature regulation below T_{lc}, and its

slope underestimates C_{mw}. Consequently, the best estimate of C_{mw}

for _Procyon lotor_ in winter is the value calculated for group A

animals with Eq. 4 (0.0172 mL O_{2}.g^{-1}.h^{-1}. deg.C^{-1}; Table 3),

and the minimum cost of thermoregulatory metabolism at any T_{a} below

T_{lc} is best estimated by substituting this value into Eq. 4 and

solving for [.H]_{r}.

Metabolic Adaptation to Climate and Distribution of the Raccoon Procyon Lotor and Other Procyonidae · 第 27 / 127 题