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The large difference in incubation time among vapor pressure difference between the egg and
bird eggs, ranging from a minimum of 11 days the microclimate of the nest is 35 torr.
to nearly 90, has aroused man's interest since
antiquity. In her critical review of the history RESULTS
of our knowledge of incubation periods, Mar- OBSERVEDINCUBATIONTIME AND
garet M. Nice (1954) wrote, "The people who EGG WEIGHT
have been concerned with incubation periods
fall into three groups: the guessers, the Heinroth (1922) plotted observed values of
incubation time against egg weight on linear
copyists, the investigators." Guessers came first
and these have been busily quoted since Aris- coordinates, while Needham (1931) and
totle for more than 20 centuries. It was not Worth (1940) used log coordinates. The lat-
until Evans (1891) and Heinroth (1922) made ter approach is mathematically more manage-
their own observations that reliable data began able and was used in our evaluation. These
to accumulate. In spite of the inaccuracies data were recalculated in order to compare
since Aristotle's time, and the many exceptions their results, based in large part upon different
which are now well recognized, there is an literature data, with our analysis. Our cor-
obvious general correlation between egg relation is based upon 475 species of birds.
Incubation periods were obtained from various
weight and incubation period. These have
been presented by Heinroth (1922), Needham sources (Groebbels 1932; Reilly 1968; Murphy
(1931), and Worth (1940) in graphic form 1936; issues of the Auk, 1940-72). Corre-
and were reinvestigated in this presentation sponding egg weights were obtained from
on the basis of newer information in the Groebbels (1969) and Schbnwetter (1960-71).
literature. It is of interest to note in retrospect These data were plotted on log coordinates
that all these correlations are essentially simi- and are shown in figure 1. A least square func-
lar, that the standard error of estimates is tion for log incubation time on log egg weight
large, and that there are many exceptions. yielded a weight power function and is shown
This merely illustrates that the many factors below for our data and authors mentioned
which determine the incubation period are above.
not understood. (1)
For these reasons we analyzed other cor- r P SE n
relates of egg size such as the gas conductance Our data
of the egg shell and particularly the water I= 12.03 WO.217 0.86 <<0.001 ?0.0922 475
loss properties of eggs, problems which Hein-
Heinroth (1922)
roth had already mentioned some 50 years ago
I = 12.09 W0.205 0.84 <<0.001 ?0.0866 194
in his classical treatise on incubation time.
On the basis of water vapor conductivity mea- Needham (1931)
surements of the egg shell previously presented I = 10 WO.240
(Ar et al. 1974), the daily weight losses of Worth (1940)
eggs during natural incubation reported by I = 12 WO.230 104
Drent (1970), and the reported incubation
periods, one is now able to derive new rela- where I = days; W = egg weight (g); r =
tionships which apply to eggs in general. correlation coefficient; P = probability coef-
These indicate that incubation time for a given ficient; SE = standard error of estimate of
egg weight is inversely porportional to the the log form of the equation; and n = number
water vapor conductance of the egg shell. of observations.
Furthermore, during natural incubation all The proportionality constants and exponents
eggs, regardless of size, lose approximately are remarkably similar among the various
18% of their initial weight and the mean water authors. We have been able to supply a cor-
relation coefficient for Heinroth's original
1 Present address: Department of Zoology, Tel-Aviv Uni-
versity, Tel-Aviv, Ramat-Aviv, Israel. analysis which is in no way improved by in-
[147] The Condor 76:147-152, 1974
10 10V10
100
2 II I I lI III I I I I I i I I jl1 1Il I I I I I I I 1 I I 1
_
.1
150 150
J_100
9
--90
so80
70
50 50
60,-,....60"
Fr o o40 0
440 ~~ 3)
20 ,• -00 .
so
o
oal? 4040
?? .,
•
- 04
e e 4
00 o
00
2D
1500: 0 0 4 0 0 001 G0 o
10 - 0 0 sor 0.
10
io o -0 .1 ogW( O2
f ? ??5
GW
ED g)
EGGWEIHT gm
1 10 10:d000 100J00
FIGURE 1. Incubation time (days) plotted against egg weight (g), n = 475. Solid line is the regression
curve, dotted line encloses the 95% confidence limits. Scale, log-log.
creasing the number of observations we have INCUBATION TIME AND WATER LOSS
added. Since the standard error of estimate OF THE EGG
is quite large, it would, therefore, suggest that
little if any refinement will be gained in the It has long been appreciated that during arti-
future by the addition of further observations. ficial and natural incubation an egg loses
Our individual data points, regression line, and weight. Not only is the weight loss under
95% confidence limits are shown in figure 1. natural conditions rather constant from day
We have tried to distinguish between al- to day (Groebbels 1932), but it can be
tricial and precocial birds, but the slight mean ascribed almost exclusively to the loss of water
difference is not statistically significant and vapor since the embryo has a typical respira-
better data are needed to establish such a fact. tory quotient near 0.72 where the exchanging
In figure 2 we have replotted the slopes of mass of 0) and CO2 molecules are equal (for
review see Drent 1973). Furthermore, this
figure 1 on a semilog plot where the incuba- water loss occurs by diffusion of water vapor
tion time is presented on a linear coordinate.
In Schbnwetter's data, we find the smallest across the egg shell and thus the total amount
transferred is determined by the pore geometry
eggs to be for two species of the family of the shell which defines the water vapor
Trochilidae, namely, 0.2-0.3 g. The largest conductance and the existing water vapor
eggs reported are those for 23 eggs of the ex- gradient between the inside of the shell and
tinct Aepyornis, which average about 9 kg, the the microclimate surrounding the egg (Wan-
largest being 12.6 kg. The longest incubation gensteen and Rahn 1970-71; Paganelli et al.
period we would predict for the Aepyornis 1971).
species is a mean value of 89 days and one These two relationships can be defined by
standard error of estimate between 72 and the following equations:
110 days. It is of interest to note that Needham
and Worth predicted a 90 and 85 day incuba- FW=M O'1I
tion period, respectively, for Aepyornis. Thus
(2)
between the largest and smallest avian egg and
we have a 45,000-fold weight difference as-
MIH20- GH APH20 (3)
sociated with a (89/11) or eightfold difference 20"
in incubation time. where W = initial weight of the egg (g)
1000r 0.74
predict that no air space develops in the egg
during incubation.
MH0=5.0 W ." For a few species the daily water loss and
'70)
(Drent : K
the water vapor conductance are known. For
100 100 example, MAH20is 450 mg day-' in the Herring
Gull (Drent 1970) and GH2O = 16 mg day-'1
torr-1 (Ar et al. 1974). Thus the mean water
vapor pressure difference in this species is
10- -o10 450/16 or 28 torr. One can estimate the water
vapor tension inside the egg shell to be 48 torr
0178 since the typical egg temperature for this spe-
GH20=0.432W cies is 37.50C. Thus the effective vapor ten-
(Aretal'73) - sion in the microclimate surrounding the egg
.
is (48-28) or 20 torr and the effective hu-
.. midity is 20/48 or 42%.
RELATIONSHIP BETWEEN INCUBATION TIME
AND WATER VAPOR CONDUCTANCE
10 100 1000
OF THE EGG
grams
EggWeight, The two generalizations, namely, the fractional
FIGURE 3. Relationship of daily water loss of eggs water loss constant and the mean water vapor
in the nest (Drent 1970) and the water vapor con- gradient, can now be used to obtain a direct
ductance of eggs (Ar et al. 1974) as a function of relationship between incubation time and
egg weight. water vapor conductance. By substituting Eq.
(3) into Eq. (2) and introducing the value
for F = 0.18 and the value of APr20 = 35 and
Substituting Eqs. (5) and (6) into the general
water vapor flux Eq. (3), we can solve for solving for I, we obtain:
the mean water vapor pressure difference, 0.18*103.W W
APH20. Thus: I= =5.2 (8)
35 GH20 GH20
(7) where 103 is introduced so that GH20 can be
15 WO.74
APHwo MHoO 0.43 W.78
= 35 torr H20 expressed in its conventional dimension of
-GH20 milligrams.
if one assumes an insignificant difference be- This equation indicates that for a given egg
tween the two exponents 0.74 and 0.78. weight, W, the incubation period is inversely
This relationship tells us that the mean proportional to the water vapor conductance,
water vapor pressure difference between the GH20, and implies a small pore area/shell
inside of the egg shell and the surrounding thickness ratio in eggs with relatively long
microclimate is 35 torr. This value is expected incubations and a large ratio for eggs with
to vary around this mean among species de- relatively short incubations. Four examples
are taken from the values reported by Ar et al.
pending upon the type of nest, the incubation
behavior, and general climatic condition. In (1974) which are shown in table 1. Eggs of
the chicken and the puffin as well as the Rhea
desert nesters, one might expect a rather large
and Emu are similar in weight but not incuba-
water vapor gradient; in hole nesters, a rela- tion time. The egg weight, conductance
tively small gradient; and among the mound values, and the reported incubation periods for
builders, which cover their eggs with decaying each species are shown as well as the predicted
material, no gradient at all and therefore no incubation based on Eq. (8). The agreement
water loss. In the latter case, one would also between reported and predicted incubation
TABLE 1. Reported and predicted incubation period for eggs having similar weight.
I I
w GH20 reported predicted
10,000
(g)
90- eggs in the nest, and incubation time as func-
_ 10000- tions of egg weight.
10,000-- 80
70 EFFECT OF ALTITUDE ON EGG WATER LOSS
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