Wald Bauer

The Consumption and Utilization of Food by Insects
G. P. WALDBAUER
Department of Entomology,
Uriiilersity of Illinois, Urbana, Illinois, US.A .
I. Introduction . 229
[I. Consumption, Growth and Utilization Indices . . 231
A. Consumption and growth . . 232
B. Digestibility and efficiency of conversion . . 233
“1. Measuring Consumption and Utilization by Weight . . 236
A. General considerations . . 236
B. The gravimetric method . . 238
C. Indirect methods using markers . . 242
IV. Food Consumption . . 246
V. Digestion and Conversion of Fresh and Dry Matter . . 250
A. Limitations of the data . . 250
B. Comparison of species . . 263
C. Comparison of foods . . 264
D. Effects of environmental factors . . 265
E. Variations with age and sex . . 267
VI. Utilization of Dietary Constituents . . 272
W. Utilization of Energy . . 278
Acknowledgement . . 282
References. . 282
I. INTRODUCTION
A great deal is known concerning the qualitative nutritional require-
ments of insects. The quantitative aspects of insect nutrition have, how-
ever, received less attention, and there have been few studies on the rates
of intake and the efficiency of food utilization. In particular, relatively
little is known concerning the intake, digestibility and efficiency of con-
version of defined diets. Quantitative work with artificial diets has
usually involved only measurements of the amount of a particular
nutrient required per unit of diet (House, 1959, 1962, 1965a). This
defines the relationship between the requirements for particular nutri-
ents (Sang, 1956, 1959), but says nothing of food intake, absolute
requirements or the efficiency of food utilization. Most measurements
229
230 G . P. WALDBAUER
of intake and utilization have been made with insects feeding on natural
foods. The information available in this area is useful, but some of it is
difficult to interpret because measurements have not been standardized,
methods have varied in accuracy and confusion has resulted from the
application of indices of utilization derived from vertebrate nutrition.
I have, therefore, included an extensive discussion of these points.
Lafon (1951) reviewed some of the earlier studies of insects other than
the silkworm Bombyx mori. Most of the numerous studies of food
utilization by Bornbyx are summarized by Legay (1957, 1958) and
Yokoyama (1963). I have omitted discussion of most of this work since
it would be repetitious in the context of this review. Recent papers on
Bombyx which are not cited by the above reviewers are as follows:
Hassanein and El Shaarawy (1962a, b), Mukaiyama and Ito (1962a, b),
Shyamala et al. (1956), Takeuchi and Kosaka (1961). Trouvelot’s (1867)
measurements of food efficiency in Antheraea polyphemus are of histori-
cal interest but too crude to be useful.
Insects as a group feed upon a remarkably diverse list of organic
substances. At the same time most species show a high degree of speci-
ficity in their choice of food. Gordon (1959) hypothesized: “Competi-
tion and natural selection gradually drive and bind each species to a
specialized food supply that it can utilize more efficiently than any of its
competitors.” Nevertheless, the qualitative nutritional requirements of
growing insects seem to be relatively uniform (Fraenkel, 1953, 1959;
House, 1962). It seems apparent that adaptive nutritional differences
must be sought on a quantitative level and that a meaningful compara-
tive nutrition of insects will not emerge until quantitative studies are
emphasized. The determination of absolute requirements for dietary
constituents depends upon the measurement of intake. Differences in
food efficiency can be demonstrated only by measuring intake and
growth. Digestibility should also be measured since it can be expected
to vary widely with different foods. The efficiency with which digested
food is used for growth will vary not only with the maintenance require-
ment for energy but also with the balance of nutrients (Gordon, 1959).
Measures of intake and utilization have played little part in the classi-
cal studies of insect nutrition although their usefulness in this connection
is obvious. For example, instances of poor growth may not be due to
the nutritional inadequacy of the diet but to a low rate of intake due to
the absence of a non-nutrient phagostimulant. Conversely, the addition
of a nutrient with phagostimulatory activity might lead to increased
growth although the nutrient is neither required nor utilized (Dadd,
1960). Sang (1959) suggested that there may be more than one optimal
CONSUMPTION A N D UTILIZATION O F FOOD 231
diet for a given species. Measures of intake and utilization can give an
indication of this, since patterns of utilization may be different although
diets are similar in their ability to support growth. For instance, low
intake might be offset by high digestibility or a high utilization of
digested food for growth or vice versa. Poor digestibility might be offset
by the efficient utilization of digested food or vice versa.
One of the major concerns of modern ecology is the elucidation of the
energetic relationships within and between communities. A knowledge
of the food efficiency of insects is thus of particular importance to
ecology since, in terms of either numbers or biomass, insects are a
major component of almost all terrestrial or fresh water communities.
Engelmann (1966) reviewed the field of terrestrial energetics and dis-
cussed its ecological significance. He also discussed the importance of
food efficiency data in ecological energetics.
11. CONSUMPTION, GROWTH A N D U T I L I Z A T I O N INDICES

An overall understanding of the utilization of a food requires answers
to the following questions: At what rate is the food eaten? How much
of the eaten food is actually digested? What part of the food is incor-
porated as body substance? These questions cannot be answered with-
out a reliable and accurate method for determining the weight of the
food ingested, the weight of the feces which correspond to the ingested
food and the weight gained by the insect during the experiment. These
three measurements are basic and necessary for the calculation of the
rate of feeding, the digestibility and the efficiency of conversion of food
to body substance. This is true whether the question is the utilization of
whole food (fresh weight), dry matter, energy, carbohydrates, nitrogen
or other nutrient constituents of the food.
Utilization data are most useful if they are reduced to terms which
allow one to compare, for example, the utilization of different foods,
utilization from instar to instar, or the effects of environmental factors
on utilization. Various indices of consumption, digestibility and effi-
ciency of conversion have been devised by the students of vertebrate
nutrition (Kleiber, 1961;Tyler, 1964). The use of these indices in insect
nutrition has been discussed by Trager (1953), Gordon (1959) and
Waldbauer (1962, 1964).
The indices mentioned below can be calculated on the basis of fresh
or dry weight, but it should be noted that fresh and dry weight indices
are not comparable since the percentage dry matter of food, feces and
insect are likely to differ.
232 G. P. W A L D B A U E R
A . CONSUMPTION A N D G R O W T H
I . Consumption index
In his classic work on accessory growth factors Hopkins (1912)
pointed out that absolute quantities cannot be used to compare the
intake of rats growing at different rates-that valid comparisons can be
made only on the basis of the rate of intake relative to the mean weight
of the animal during the feeding period. The consumption index (C.I.)
is, therefore, calculated as :
r
C.I. =
TA
_f_
F = fresh or dry weight of food eaten

T = duration of feeding period (days)
A = mean fresh or dry weight of animal during feeding period.
The mean weight of the animal is most accurately calculated from the
area under its growth curve as determined by integration or direct
measurement. A weighted average of daily weights will give an almost
identical value if the growth curve is smooth (Waldbauer, 1964).
Other indices of consumption have been proposed. Lafon (1951) used
mg of food eaten per g of insect (final weight) in thirty days. The useful-
ness of this index is difficult to discern since it neglects the initial weight
of the insect. For instance, an insect which eats 500 mg as it grows from
10 to 100 mg is obviously eating at a greater rate relative to its weight
than an insect which eats the same quantity in the same time as it grows
from 50 to 100 mg. Legay (1957) proposed mg of food eaten per g of
insect (initial weight) per day. In this case two insects of the same initial
weight which eat the same quantity of food in the same time will have
identical indices even if one grows more than the other. Smith (1959)
used weight of food eaten divided by weight gained. This yields an
infinitely large index if the insect eats but does not grow, a dramatic
expression of the failure to grow but not a useful measure of feeding
rate.
See Section IV for further discussion of feeding rates.
2. Growth rate
The reZatiue growth rate (G.R.) is calculated as:
G
G.R. = -
TA
CONSUMPTION A N D U T I L I Z A T I O N OF FOOD 233
G = fresh or dry weight gain of animal during feeding period
T = duration of feeding period (days)
A = mean fresh or dry weight of animal during the feeding period.
Workers who measured efficiency of utilization in insects usually did
not compute true growth rates (see, however, Waldbauer, 1964).
B . DIGESTIBILITY A N D EFFICIENCY OF CONVERSION
1. Conversion of ingested food

The eficiency of conversion of ingested food to body substance (E.C.I.)
is calculated as :
wt gained
E.C.I. = x 100
wt food ingested
or
G.R.
E.C.I. = -
C.I.
The E.C.I. is an over-all measure of an insect’s ability to utilize for
growth the food which it ingests. The E.C.I. will vary with both the
digestibility of a food and the proportional amounts of the digestible
portion of that food which are, on the one hand, converted to body
substance and, on the other hand, metabolized for energy to maintain
life. Thus the E.C.I. will rise and fall with the A.D. (approximate
digestibility) and the E.C.D. (efficiency of conversion of digested food
to body substance).
2. Digestibility
The approximate digestibility (A.D.) is calculated as :
wt of food ingested-wt of feces
A.D. =
wt of food ingested
Many authors including House (1965a), Trager (1953) and Waldbauer
(1964) referred to this measure as the “coefficient of digestibility”. How-
ever, this is misleading since, as will be discussed below, the difference
between the weight of ingested food and the weight of the feces does not
represent the amount actually digested. It should be noted that approxi-
mate digestibility as here defined for insects differs from apparent
digestibility as defined for mammals.
Although mammals pass urine and feces separately, their feces do not,
strictly speaking, consist only of undigested food nor do they contain all
234 G . P. W A L D B A U E R
of the undigested food. The gases produced in digestion, mainly C 0 2

and CH4, are lost. Mammals on a nitrogen-free diet continue to pass
nitrogen in the feces. This metabolic fecal nitrogen presumably comes
from intestinal secretions and the debris of gut cells. Thus, with mam-
mals the true digestibility is not obtained by subtracting the weight of
the feces from the weight of the ingested food. The value obtained is
called the apparent digestibility and is considered to be a reasonable
approximation of the true digestibility (Kleiber, 1961; Mitchell, 1964;
Tyler, 1964). The feces of insects are certainly not free of fecal metabolic
products; the peritrophic membrane is an obvious example.
The presence of the urine in the feces further complicates the measure-
ment of digestibility in insects. The difference between the weight of
food ingested and the weight of the feces actually represents the food
which is stored or metabolized less metabolic wastes discharged in the
urine or as fecal metabolic products. Thus, the above formula yields
what I have called approximate digestibility, a value which is always
lower than the corresponding apparent digestibility. The magnitude of
the difference depends upon the relative amount of urine in the feces.
Birds mix the urine and feces in the cloaca and thus present a similar
problem (Kendeigh, 1949; Mitchell, 1964). Kendeigh refers to the
energy in the ingested food less the energy in the excrement as the
metabolized energy.
It can be argued that the use of “digestibility”-even in the sense of
“ approximate digestibility”-is inappropriate unless one accounts for
at least the urine content of the feces. However, the available alterna-
tives, “retention” (Evans, 1939b) and “utilization” (Kasting and
McGinnis, 1959; Hirano and Ishii, 1962) are also inexact. Furthermore,
“utilization” should be reserved as a general term which includes
digestion, metabolism and conversion to body substance.
An approximation of apparent digestibility can be obtained if the
estimated urine content is subtracted from the weight of the feces. I will
refer to the value thus obtained as the coeficient ofapparent digestibility
(C.A.D.).
. Determination of the C.A.D. may be an unnecessary refinement with
most insects which eat foods with a moderate protein content. Their
feces will probably contain relatively little urine, and thus the difference
between the A.D. and the C.A.D. is likely to be very small. The major
component of the urine of terrestrial insects is uric acid (cf. Patton,
1953; Wigglesworth, 1965). Analyses of the feces of phytophagous
insects indicate a low uric acid content. It ranges from 051% for the 1st
instar down to 0.24% for the 5th instar in the dried feces of Bombyx
CONSUMPTION A N D UTILIZATION OF FOOD 235
mori (Hiratsuka, 1920). Subtracting the uric acid content from the feces
results in a negligible increase of the A.D. It is raised from 46.0% to
46.4% for the 1st instar and from 36.5% to 36.7% for the fifth instar
(calculated from data of Hiratsuka, 1920). The dried feces of other leaf-
feeding insects are also low in uric acid, 0.62% for the 4th instar and
0.52% for the 5th instar of Neodiprion sertifer (Janda, 1961), and about
4% for Melanoplus bivittatus (Brown, 1937a). A similar correction for
the 4% uric acid content of the feces of Tenebrio molitor larvae raises
the A.D. from 46.3% to 485% (Evans and Goodliffe, 1939). Gupta and
Sinha (1960) found a higher concentration of uric acid in the feces of the
adults of stored grain beetles, about 12% of the dry weight for Sitophilus
granarius, 18% for Tribolium confusum and almost 20% for Cryptolestes
ferrugineus.
The feces of insects which eat foods high in protein are likely to con-
tain a relatively greater percentage of urine. The approximate digesti-
bility may, therefore, differ greatly from the coefficient of apparent
digestibility. The larvae of the webbing clothes moth, Tineola bisselliella,
eat hair and other substances which consist largely of keratin. Their
dried feces contain about 28% uric acid plus far smaller amounts of
ammonia and urea (Hollande and Cordebard, 1926). Using this figure
to correct Titschack’s (1925) data for Tineola (Table V) raises the A.D.
of 29% to a C.A.D. of approximately 56%.
3. Conversion of digested food

The eficiency with which digested food is converted to body substance
(E.C.D.) is calculated as:
wt gained
E.C.D. = x 100
wt of food ingested - wt of feces
E.C.D. will decrease as the proportion of digested food metabolized
for energy increases. Thus the E.C.D. is affected by factors which influ-
ence the amount of energy devoted to the maintenance of physiological
functions or the support of activity. E.C.D. is not directly dependent
upon digestibility, but it does vary with the level of nutrient intake.
Since the maintenance requirement remains more or less constant, the
proportion of food available for growth will decrease as intake decreases.
E.C.D. also varies with the nutritional value of the food. For example,
Hopkins (1912) demonstrated that a vitamin deficiency caused rats to
waste an enormous amount of absorbed food (see Gordon’s (1959) lucid
discussion of the significance of food efficiency).
111. MEASURING A N D UTILIZATION

CONSUMPTION B Y WEIGHT
A . GENERAL CONSIDERATIONS
The selection of the period during which utilization is to be measured
is critical. An arbitrarily defined period, a specified length of time not
marked by physiological events, offers the advantage of convenience.
However, data obtained on this basis are of limited value since, as will
be shown below, utilization varies quantitatively from instar to instar
and within instars. Physiologically defined periods are less convenient,
but are more likely to yield reproducible results which can be validly
compared with the results of other experiments.
A physiologically defined period may include the entire life cycle, an
entire stage or one or more instars. In any case its beginning and end are
marked by physiologically significant events. It may begin with hatch-
ing or the completion of a molt. Its end may be marked by the premolt
condition, completion of a molt or an event in the life of the adult. The
use of physiologically defined periods presents some inconveniences. It
may be difficult to obtain enough newly hatched or molted insects to
begin an experiment. Growth rates may vary, necessitating frequent
inspections to fix the end of the period for each individual.
Physiologically defined periods were used by Crowell (1 941), Hirat-
suka (1920), So0 Hoo and Fraenkel (1966), Waldbauer (1964) and
others. Some workers used mixtures of instars or failed to specify instar
or the length of the period. Others, for instance Evans (1939b) and
Chauvin (1946), used arbitrary periods of time within specified instars.
Evans (1939b) stated that he used larvae of different physiological ages
within the same instar.
Accuracy demands the collection of all feces derived from the food
eaten during an experiment. Residual food which may be in the gut at the
beginning or the end of an experiment can cause an error in both weight
gain and the weight of feces. The gut contents of feeding early 4th
instar Protoparce sexta is about 9.3% of the fresh body weight (Wald-
bauer, 1962). McCay (1938) found that residual food in the gut of feed-
ing Blattella germanica amounts to less than 5% of the dry body weight.
The use of physiologically defined periods simplifies quantitative collec-
tion of the feces since the gut will be more or less empty just before or
after a molt. It seems that most insects empty the gut before each molt.
Indeed food could be retained only in the midgut since the linings of
both fore- and hindgut are molted. Protoparce larvae do not retain a
significant residue of food through the molt (Waldbauer, 1964). How-
ever, So0 Hoo (1962) states that considerable food may be retained in
the midgut of Prodenia eridania after a molt.
The larvae of Protoparce empty the gut during the first few hours of
the premolt period (Waldbauer, 1964). The time required to empty the
gut after cessation of feeding may be related to the speed with which
food passes through the gut during feeding. House (1965a) summarized
passage times which range from 25min for adults of the cucujid
Oryzaephilus surinamensis to 2-73 hr for Prodenia larvae and 28-32 hr
for adult females of Aedes aegypti.
With farm animals the feces derived from the experimental food are
often identified by feeding an indigestible marker at the beginning and
end of the experiment. Only those feces which appear between the two
markers are collected (Kleiber, 1961). Foods which produce feces of
different colors were used to the same end by Phillipson (1960) in his
study of the food consumption of a phalangid. Similar methods have
apparently not been used with insects although Crowell (1943) and
Husain et al. (1946) used indigestible markers to measure the rate at
which food passes through the gut. Chauvin (1946) tried to eliminate
residual food by starving adult Orthoptera for 24 hr before the experi-
ment and until they stopped passing feces after the experiment. This
procedure may not injure grasshoppers, but insects which feed almost
continuously (i.e. aphids or many lepidopterouslarvae) may be seriously
stressed by a short period of starvation. Furthermore, Waldbauer (1964)
found that starved larvae retained considerably more residual food than
newly molted ones. Evans (1939b) devised an elaborate indirect method
using four groups of larvae. At the beginning of the experiment the first
group was sacrificed and dried to a constant weight. The second group
was starved and their feces dried and weighed to provide an “excreta
equivalent”. The two remgiling groups were provided with food. At
the end of the feeding period the experimental group was dried and
weighed. The fourth group was then starved and another “excreta
equivalent’’ obtained. The difference between the two “excreta equi-
valents” was added to the weight of the feces of the experimental group
and subtracted from their dry weight gain. Evans’ (1939b) method
avoids long and possibly deleterious starvation, but is certainly tedious
and probably less accurate than the use of a physiologically defined
experimental period.
The molting of insects presents a special problem in the determination
of the E.C.I. and the E.C.D. since the insect reaches a maximum weight
in each instar and loses weight during the molt. Both the molted cuticle
and the energy used during the molt contribute to this loss. Fifth instar
Protoparce which attained a maximum fresh weight of 7.14 g (after

emptying the gut) weighed only 3.94 g after molting to the pupa, a loss
of about 45% (Waldbaucr, 1962). Thus a net weight gain lower than the
gross or total weight gain is obtained if the experimental period includes
one or more molts and if gain is taken as the difference between the
weight of the insect at the end and the beginning of the experiment. Use
of the net weight gain yields a net E.C.I. or E.C.D., a considerably
lower figure than the gross E.C.I. or E.C.D. which include the weight
lost at each molt.
Some workers used groups of insects rather than individuals. Hirat-
suka (1920) began his experiments with four groups of lo00 Bombyx
mori larvae. Evans and Goodliffe (1939) used a single group of over
2000 Tenebrio molitor larvae. However, the use of groups may be com-
plicated by mortality, cannibalism or the eating of dead individuals.
With many insects, especially those which feed on powdered diets, the
likelihood that they will eat feces is increased. Hiratsuka (1920) mini-
mized the problem of mortality by removing all sick larvae each day
and replacing them with healthy larvae of similar size. Evans and
Goodliffe (1939) did not mention the problem of mortality. Crowell
(1941), Evans (1939a, b), Nagy (1953), Balogh and Gere (1953) and
Davey (1954) used smaller groups. Crowding is likely to affect the rates
of feeding and growth (see Section VD).
The size of the container in which the experiment is carried out may
be of importance. Fewkes (1960) found that the 1st and 2nd instars of
the hemipterous predator Stalia major had a lower E.C.I. than the
latter instars. He felt that this may have been due to the use of similar
sized cages for all instars. The ratio of search area to size of predator
was higher for the early instars. Thus they required proportionally more
energy to obtain each meal.
B . THE GRAVIMETRIC METHOD
Most workers have used the classical gravimetric method to measure
the utilization of food. Other methods will be discussed in Section IIIc.
In essence the gravimetric method involves the following procedure:
1. The insect’s weight gain is found by subtracting its weight at the
beginning of a feeding period from its weight at the end. 2. Food eaten
is determined by subtracting the weight of uneaten food from the weight
of the food provided. 3. All feces are separated from the uneaten food
and weighed.
If the experimental period includes a molt the difference between
initial and final weights yields the net weight gain of the insect. Gross
gain is then obtained by adding to net gain the weight lost during each
molt. This loss is determined by subtracting the weight of the newly
molted insect from the maximum weight attained in the previous instar.
Some of the major difficulties encountered in the determination of
weight gain should be mentioned. The presence of residual food in the
gut can cause a considerable error as discussed above. The best solution
is to begin the experiment with newly molted larvae whose guts are
naturally empty, and to end with either premolt or newly molted larvae
whose guts are again naturally empty. An additional error is avoided by
weighing individuals as soon as possible after they have molted or
entered the premolt stage. Premolt Protoparce which are about to under-
go a larval molt are inactive and lose only about 0.25% of their fresh
weight per hour. Newly molted larvae are more active and can be
expected to lose more weight. Larvae preparing for the pupal molt are
extremely active and lose about 1.3% of their fresh weight per hour
(Waldbauer, 1960). If an experiment need include only one instar the
penultimate is the most convenient choice since it eats fairly large
quantities and the weight lost during the premolt is at a minimum. The
weight of silk or other products must be considered if gross efficiency is
to be calculated.
It is, of course, impossible to measure dry weight directly at the
beginning of an experiment. An estimate can be calculated from the
mean per cent dry matter of an aliquot of similar larvae dried to con-
stant weight at 100°C. It is convenient to kill the insects first by placing
them in a freezer for a short time. At the end of the experiment dry
weight may be determined either directly or by means of another
aliquot.
A d libitum feeding is convenient and natural, and has generally been
used with insects in preference to some form of controlled feeding.
Taking the weight of the food eaten as the difference between weight
of the food provided and weight of the uneaten food poses some prob-
lems. If fresh weights are used it is necessary to control for a loss or
gain of water by the food. Water loss can be minimized by maintaining
a humid atmosphere. Evans (1939a, b) estimated the loss from leaves
provided as food from the loss of similar leaves kept without insects in
containers similar to those used in his experiments. He calculated the
“natural loss” of the food as:
% wt loss of aliquot x wt food introduced2 + wt uneaten food

A more precise estimate is obtained if the correction factor is calculated
240 G. P. WALDBAUER
both as the ratio of loss to the initial weight of the aliquot (a) and as the
ratio of loss to the final weight of the aliquot (6). Then:
corrected wt food eaten
W = wt of food introduced
=
[1 - -4 [W - ( L + bL)]
L = wt of uneaten food.
In this calculation the full correction is applied to the uneaten food
because it lost weight during the entire experimental period. One-half
the correction is applied to the eaten food since it is assumed that it was
being eaten at a constant rate. This calculation differs slightly from one
suggested by Waldbauer (1962). The use of an aliquot to estimate
“natural loss” in this way involves at least two additional assumptions:
I . The % weight loss of the food is independent of the amount of food.
2. Feeding itself does not increase “natural loss” by constantly provid-
ing a freshly cut surface.
The absolute amount of “natural loss ” and, thus, the relative size of
the error will increase with the amount of uneaten food which remains
at the end of the experiment. If excess food is held to a minimum
“natural loss” will often be negligible (Carne, 1966 and Waldbauer,
1962). However, Waldbauer (1962) found that excised leaves of some
plants lose as much as 4% of their initial weight in 24 hr.
The dry weight of foods which cannot be dried before feeding must be
estimated from the % dry matter of an aliquot. Some foods, particu-
larly leaves, demand judicious selection of the aliquot. Chauvin (1946)
found that variations in the dry matter content of leaves necessitated
the cutting of an aliquot from each leaf used. Brennikre et al. (1949),
Waldbauer (1964) and So0 Hoo and Fraenkel(l966) found that greater
precision is obtained by cutting the leaves into two symmetrical por-
tions along the midrib, using one portion as food and the other as the
aliquot (Table I). Hiratsuka (1920), Evans (1939a, b), Smith (1959) and
others used whole leaves as aliquots. This probably had little effect on
the accuracy of Hiratsuka’s (1920) results since he used very large
quantities of leaves.
The dry weight loss of excised leaves by metabolism is probably
negligible. It can be minimized by frequent changes of food and the
immediate drying of uneaten food. The potential error from this source
could be further minimized by estimating the dry weight of eaten food
from the mean of the % dry matter of the aliquot and the % dry matter
of the uneaten food.
The small but inevitable discrepancy between the % dry matter
TABLE I
A comparison of the use of whole leaves and asymmetrically and symmetrically
cut leaf pieces as aliquots for the estimation of % dry matter of whole leavesa
Range of % dry Mean difference in % dry matter of:

matter of whole apical and basal right and left
Plant leaves halves of leaves portions of leaves
Lycopersicon
esculentum 12’7-16.6 1.02 0.16
Solanum fuberosum 11.5-17.2 1-32 0.58
Taraxacum oficinafe 13.6-1 7.0 2-01 0.14
Arctium minus 17.4-245 2.04 0.26
* From Waldbauer (1964).
determined from the aliquot and the actual % dry matter of the food
causes an error in the estimated dry weight of the food provided. The
dry weight of uneaten food is not subject to this error since it is measured
directly. The error is passed on when food eaten is calculated by sub-
tracting weight of uneaten food from the weight of food provided. The
absolute size of the resulting error in weight of food eaten will increase
with the quantity of food introduced, while its relative size will increase
with the proportion of uneaten food. Adding an excess equal to the
quantity eaten will thus double the error.
The food intake of blood-sucking insects can be conveniently deter-
mined by weighing the insect immediately before and after a meal.
Buxton (1930) and Friend et al. (1965) used this method with Rhodnius
prolixus. Titschack (1930) and Johnson (1937 and 1960) used it with
Cimex lectulurius. Titschack (1930) pointed out that Cimex passes a
drop of excrement immediately after feeding.
Fewkes (1960) and Evans (1962) measured the food intake of pre-
daceous Hemiptera by supplying weighed prey which were reweighed
after the predators had fed.
The attempts to measure the rate of sap intake by aphids were re-
viewed by Auclair (1963). Robinson (1961) found that weighing pre-
viously starved aphids before and after feeding did not give reliable
results. This was probably because they began excreting honeydew as
soon as they resumed feeding. Experiments with reproducing adults
were further complicated by the birth of nymphs. Mittler (1957) tried to
estimate sap intake from the amount of sap which exudes from em-
bedded stylets severed from feeding aphids. However, the rate of
242 G . P. W A L D B A U E R
exudation was sometimes somewhat less than the rate of honeydew

excretion by comparable aphids. Apparently the turgor pressure of the
plant is not solely responsible for the flow of sap into the aphid’s ali-
mentary canal.
The dry weight of the feces can be measured directly. Feces should be
collected frequently and dried as soon as possible to avoid decomposi-
tion. Brown (1930) found that after 24 hr there had been a marked
change in the carbohydrate composition of the moist feces of Automeris
io. He thought that the change was due largely to the growth of mold.
The fresh weight of feces may be difficult to determine because of a
loss or gain of water. The feces of leaf-feeding insects are likely to lose
water rapidly. Those of the larvae of Automeris lose about 26% of their
fresh weight in 24 hr (Brown, 1930). Evans (1939a, b) minimized this
loss by conducting experiments in air-tight cans and collecting the feces
each hour.
Legay (1953, 1957) devised an ingenious but tedious gravimetric
method of measuring food intake and excretion in fresh weight. He
placed a Bornbyx mori larva on the pan of a beam balance, noted its
initial weight and gave it a leaf to eat. Every 15 min he removed the leaf
and noted the larva’s weight gain. The weight of freshly passed feces
was taken by noting total weight before and after brushing the pellets
from the pan. The weight of food eaten was taken as the weight gained
by the larva between successive weighings plus the weight of feces
passed. Legay (1957) pointed out that his method does not account for
the weight of water vapor or carbon dioxide lost by the larvae. Legay
felt that this error is small but significant, and suggested that it could
be estimated by measuring the weight loss of starving larvae. The error
is not small. Recalculating the data of So0 Hoo and Fraenkel(l966) for
Bornbyx on the basis of fresh weights (using Legay’s (1957) figures for
water content of feces) shows that of the food eaten by the 4th instar
29% is excreted as feces, 48% is used for growth and 23% is metabolized
and thus lost in the form of water and carbon dioxide. Furthermore,
frequent disturbance of the larvae may seriously affect their feeding
rate. Legay’s (1957) method is certainly useful for determining the fresh
weight of feces, but is of doubtful value for measuring intake.
C . I N D I R E C T M E T H O D S USING M A R K E R S
Indigestible markers have been used to measure the utilization of food
by vertebrates, especially farm animals, for the last forty years (see Elam
et al., 1962; Kleiber, 1961;Maynard, 1951;Smart et al., 1954). The most
frequently used method involves the addition of a known concentration
of chromic oxide to the diet and the determination of the concentration
of chromic oxide in the corresponding feces. The C.A.D. or A.D. can
then be determined without quantitative collection of the feces or
ineasurement of the food eaten:
M F = concentration of marker in food

M E = concentration of marker in excreta.
If either the weight bf feces or the weight of food eaten are known it is
possible to calculate:
wt food eaten = 5
MF
x wt feces
or
wt feces =
MF x wt food eaten
-
ME
McGinnis and Kasting (1964a) developed a sensitive colorimetric
analysis for chromic oxide which made it possible to use the marker
technique with individual insects. The technique was first used to
measure the A.D. of insects, larvae of Agrotis orthogonia, by McGinnis
and Kasting (1964b). Later they compared the A.D.3 and dry weight
food consumption of nymphs of Melanoplus bivittatus and larvae of
Agrotis as measured by the chromic oxide and gravimetric methods
(McGinnis and Kasting, 1964~).They claimed that the two methods
gave more or less comparable results. However, in three of six tests the
chromic oxide method gave A.D.’s considerably lower than correspond-
ing A.D.’s obtained gravimetrically. They concluded that the gravi-
metric method had overestimated the A.D., arguing as follows: 1. One
of their diets and the resulting feces were crumbly, making it difficult to
separate uneaten food and feces for gravimetric determination. Com-
plete separation is not required for determination of the A.D. with the
chromic oxide method. 2. Variations in the % dry matter of aliquots led
to an error in the calculated dry weight of the food introduced. They
had, however, introduced almost four times as much food as the insects
ate, greatly increasing the relative size of this error (see discussion
above). They also found that the chromic oxide method gave less
variable A.D.’s than the gravimetric method. However, with the gravi-
metric method Waldbauer (1964) obtained A.D.’s with standard devia-
tions of about the same size as those obtained by McGinnis and Kasting
(1964b, c) with the chromic oxide method.
244 G . P . WALDBAUER
The chromic oxide technique was subsequently used by McMillian et

al. (1966) to compare the digestion by Heliothis zea and Spodoptera
frugiperdu of various lyophilized plant parts incorporated in an agar
diet fortified with vitamins. Larvae on these diets gained less weight than
larvae on the usual laboratory diet. The authors suggest that the
chromic oxide may have deterred feeding.
McGinnis and Kasting (1964~)stated that feces need not be collected
quantitatively in order to determine the A.D. In fact they did collect
feces quantitatively and analyzed all of the feces of each insect for
chromic oxide. It should be noted that a sampling error may result if
only a part of the feces are used for analysis. Elam et al. (1962) reported
that they and others observed that the concentration of chromic oxide
in the feces of sheep, goats and cattle appears to vary with time. As yet
we have no idea of whether or not sampling errors will be a serious
problem in using the chromic oxide method with insects.
As pointed out by McGinnis and Kasting (1964b), a marker used to
measure food utilization should meet the following criteria: 1. It should
not affect feeding or be toxic at the concentrations used. 2. It should not
be altered by or absorbed from the gut. 3. It should be possible to
distribute the marker uniformly in the food. 4. An accurate and rela-
tively rapid analysis for the marker should be available.
According to McGinnis and Kasting (1964~)the chromic oxide
method is more rapid and practical and avoids some of the error which
may result from use of the gravimetric method. The gravimetric method
is laborious but, as shown above (Section IIIB), error can be minimized
or reduced to insignificance. McGinnis and Kasting (1964~)pointed out
that for determination of food consumption both methods require the
quantitative separation of food and feces, often a difficult procedure.
With the chromic oxide method uneaten food need not be quantitatively
collected, thus allowing greater flexibility in the choice of methods of
separation.
The chromic oxide method is probably usable only with foods which
can be ground and mixed to assure even distribution of the marker. The
utilization of whole leaves could be measured if it were possible to distri-
bute the marker evenly on the surface of a leaf. However, the amount of
chromic oxide ingested would depend upon the area rather than upon
the weight of leaf eaten. Thus an accurate determination is possible only
if the relationship between leaf area and weight is known and constant.
A naturally occurring constituent of the food would be the ideal
marker. The constituent chosen should be evenly distributed in the food,
not vary greatly in concentration, and be amenable to rapid and accurate
analysis. Ideally it should not be absorbed from the insect’s gut. Elam
e l aZ. (1962) found that using lignin as a marker with sheep gave a low
C.A.D. because all of the lignin in the food was not recovered in the
feces. They state that while some workers found varying degrees of
lignin digestibility others found lignin to be a satisfactory marker.
Brown (1 930) used the starch in leaves as a marker for the determination
of A.D. in Automeris larvae. The results were similar to those he ob-
tained with a gravimetric method which was probably inaccurate.
Chromogens, naturally occurring plant pigments, have been used as
markers to measure the digestibility of dried, ground plant material by
grasshoppers (Heinrichs and Pruess, 1966). Their method is an adapta-
tion of one devised by Reid et al. (1950) for use with dairy cows. Hein-
richs and Pruess (1966) claimed a general correlation between the results
of the chromogen-ratio and gravimetric methods, although the results
differed widely in four of twenty-one tests with different plants or plant
parts. They did not describe their gravimetric method. Further work on
the development of methods using naturally occurring markers would
certainly be worthwhile. Such methods should allow one to measure
utilization by phytophagous insects living on their food plants in the
field. Perhaps similar studies would be possible with predators or blood-
feeders.
Radioactive tracers have also been used to estimate food consump-
tion. Crossley (1963a) stated that when an insect reaches a steady-state
concentration of a radioisotope through feeding on tagged food, then
the rate of intake is equal to the rate of “biological elimination” of the
radioisotope (see also Crossley, 1963b, 1966). Crossley (1963a) used
cesium-137 to estimate the intake of Chrysomelu knabi feeding on SaZix
nigra growing in a former disposal area contaminated with the radio-
isotope. Cesium-137 estimates of the daily food consumption of 3rd
instar larvae in the field agreed closely with the results of direct labora-
tory measurements.
S ~ c r o s e - U - ~or
~ Ccell~lose-U-~~C were used as markers to estimate
food consumption by 1st and 5th instar Agrotis (Kasting and McGinnis,
1965). At the end of the feeding period the insects, their feces and the
CO, they had expired were measured for radioactivity. The use of
s ~ c r o s e - U - ~with
~ C 5th instar larvae gave results which were generally
higher than those obtained with either the gravimetric or the chromic
oxide method. Kasting and McGinnis (1965) suggested that the larvae
may have extracted moisture from the diet independently of dry matter,
thus increasing their ingestion of the soluble s~crose-U-~~C. The use of
insoluble cell~lose-U-~~C gave results which agreed closely with those
obtained by the chromic oxide method. Larvae fed a diet containing

~ellulose-U-~~C produced an amount of 14C02equivalent to less than
1% of the 14Cingested. No measurable radioactivity remained in their
bodies. The 14C02 produced by larvae fed a diet containing sucrose-
UJ4C was equivalent to about 10% of total consumption. A large
amount of radioactivity remained in their bodies.
Krishna and Saxena (1962) found that the gravimetric method could
not be used to estimate food consumption by Tribolium custuneum or
Trogoderma grunarium because the sticky feces and the flour diet could
not be completely separated. They then devised an indirect method
based upon their assertion that ethanolic extracts of larval feces
do not contain sugars which react with anthrone. Thus, they used
colorimetric determinations with anthrone to estimate the weight of
uneaten flour in an inseparable mixture of flour and feces by the
following calculation :
wt uneaten flour =
S2
- x wt of flour introduced
s
1
S1= total sugars in ethanolic extract of a sample of flour

S, = total sugars in ethanolic extract of a sample of the mixture of
feces and uneaten flour.
Any food constituent which is completely absorbed or completely
altered might be similarly used to determine the proportions of feces and
food in an inseparable mixture.
I V . FOODCONSUMPTION
The dry weight-fresh weight consumption index (C.I.) is calculated
from the dry weight of the food eaten and the fresh weight of the insect.
It is of nutritional interest since it measures the rate at which nutrients
enter the digestive system. The fresh weight C.I. (calculated from fresh
weight of food and insect) is probably the better measure of the be-
havioral response to the food. Physiological and behavioral responses
are not independent of each other and are certainly not totally separable.
However, it seems reasonable to assume that the feeding rate is limited
by the response to the bulk, water content and other physical and
chemical properties of whole fresh food, and that the rate of dry matter
intake is largely a function of this response.
So0 Hoo and Fraenkel(l966) and Waldbauer (1964) found that dry
weight C.I.’s (calculated from dry weight of food and insect) were
C O N S U M P T I O N A N D U T I L I Z A T I O N O F FOOD 247
always higher than the corresponding fresh weight C.I.'s because the
insects in question contained a lower percentage of dry matter than their
food. The rank order of either dry weight-fresh weight or dry weight
C.I.'s is not necessarily the same as the rank order of corresponding
fresh weight C.I.'s. This is, of course, due to differences in the % dry
matter of the food. Thus dry weight intake may be higher on one food
than another although fresh weight intake is lower (Table 11).
TABLE
I1
Summary of Consumption Indices (C.I.)
% dry C.I. calculated from :

Food plant matter of Fresh wt food Dry wt food Dry wt food
leaves Fresh wt Fresh wt Dry wt
larva" larvab larva"
Ly copersicon
escuIentum 17.28 1*87 0.33 2.92
Lycopersicon
esculentum 15439 2.22 0.35 3.30
Lycopersicon
esculentum 15.87 2.13 0.34 3.08
Lycopersicon
esculentum 15.93 1 a87 0.30 2.67
Solanum tuberosiim 13.60 1.96 0.26 2.40
S. dulcamara 19.75 1.84 0.37 3.22
Taraxacum
oficinalec 21.12 1 *66 0.34 3.14
Arctium minusC 19.79 I 44 0.28 2.74
Verbascum
thapsus". 25.81 1 *28 0-33 3.14
* From Waldbauer (1964). Unpublished data of Waldbauer. Not normally eaten;

accepted only after maxillectomy. Supports only slow growth.
So0 Hoo and Fraenkel(l966) presented a number of dry weight C.I.'s

for 5th instar Prodenia. The daily dry matter intake on plants which
supported good growth was from 1.2 to 2.4 times the dry body weight
of the feeder. On plants which did not support growth or supported
little growth the daily dry matter intake ranged from 1.0 to 2.2 times
the dry body weight. Thus in some cases the rate of intake may be
greater on a nutritionally inadequate diet than on an adequate diet (see
also Table 11).
The rate of food intake by Celerio euphorbiae larvae appears to increase
with decreases in the nutrient level of the diet. House (1965b) used four
248 C. P. W A L D B A U E R
diets which contained the same amounts of cellulose and agar, decreas-
ing amounts of the same balance of nutrients (85%, 70% and 50% of
the highest nutrient content), and correspondingly increasing amounts
of water, approximately 87%’ 89%’ 90% and 92% of the respective
diets (the latter calculated from House’s data). Total fresh weight con-
sumption increased with the dilution of the nutrients. Thus the larvae
partly compensated for a low nutrient concentration. Approximate
mean C.I.’s calculated from House’s (1965b) data show this nicely.
Fresh weight C.I.’s were 1.4, 1.5, 1-6 and 1.8 in order of decreasing
nutrient concentration. The corresponding dry-fresh weight C.I.’s were
0.20, 0.20, 0.18 and 0.16. McGinnis and Kasting (1966) found that
grasshoppers more than doubled their intake when the nutrient content
of the diet was halved by replacing half of the lyophilized wheat sprouts
with cellulose. In this case the % H 2 0 in the diet remained the same.
House’s (1965b) comparison of a balanced diet with a nutritionally
imbalanced diet is not so clear. Total fresh and dry weight intake were
lower on the imbalanced diet, but the differences cannot be shown to be
statistically significant. Approximate mean C.I.’s calculated from
House’s data suggest that the rate of fresh weight intake of the im-
balanced diet (0.86) was lower than the rate of fresh weight intake of the
balanced diet (0.95). The rates of dry matter intake are identical (0.14)’
presumably because of the greater dry matter content of the imbalanced
diet, 15.8% versus 12.7% in the balanced diet (calculated from House’s
data).
These observations raise interesting questions concerning the be-
havioral and physiological control of the feeding rate. Is there a direct
behavioral response to increased succulence or dilute flavor? How do
the physiological effects of a diet influence the rate of feeding? Can
balanced and imbalanced diets taste noticeably different to insects? If
so, can an imbalanced diet be avoided because its taste comes to be
associated with physiological disturbances by conditioning? The reader
is referred to the papers of Garcia et al. (1967) and Revusky and Bedarf
(1967) for related discussions.
There appears to be a negative correlation between the rate of dry
matter intake and the efficiency of conversion of ingested or digested
food by Prodeniu larvae (So0 Hoo and Fraenkel, 1966). House’s (1965b)
data seem to suggest a similar relationship for the larvae fed diets with
varying nutrient concentrations. As the mean dry-fresh weight C.I.’s
decreased from 0.20 to 0.16 the corresponding mean E.C.T.’s increased
from 18 to 22 (calculated from House’s data).
Table 111 shows the proportional food intake of the larval or nymphal
C O N S U M P T I O N A N D U T I L I Z A T I O N OF FOOD 249
111
TABLE
Per cent eaten by each instar of the total quantity of food eaten during the entire larval
or nymphal stage
Insect, food and authority O; of total consumption by instars

I It I11 IV v VI VII
_. -
~- - .. -. ~
Agrotis orthogonia
Triticum aestivum sprouts 0.2 I 0.42 2.3 8.7 31.6 56.8
Triticuni durum sprouts 0.15 0.48 3.1 9.1 32.9 54.2
Kasting and McGinnis (1959)"
Agrotis ypsilon
Corn foliage 0.06 0.18 0.77 2.6 10.4 86.0'' -
Satterthwait ( 1 933)' 0.04 0.1 I 0.52 1.6 4.4 19.6 73.7"
Bonibyx mori
Morus leavesh 0.08 046 2.6 12-0 84-8 -
Hiratsuka (1920)
Caraicsius morosils
Hedera helix leaves" 1.9 3.3 6.7 11.6 24.9 51.5
Titschack (1924)
Paropsis atomaria
Eucalyptus shoots" 1.8 2.2 19.4 75.5 - -
Came (1966)
Pieris brassicae
Cabbage leaves 0.12 0.56 2-1 11.5 85.7 -
David and Gardiner (1962)"
Protoparce sexta
Tobacco leaves" 0.08 0.53 1.9 10.5 86.4 -
Wolcott (1 937)
Rhodnius prolixirs
Rabbit blood" I *2 3.3 10.3 25.0 60.1 -
Buxton (1930)
Schistocerca gregariu
Ficus leavesa I .9 4.4 11.1 20.0 62.7 -
Husain et al. (1946)
Stalia major
Carpophilus larvae' 3.8 7.2 12.2 26.8 50.1 -
Fewkes (1960)
~~ ~ ~~~~ ~
.' Measured in fresh weight. Measured in dry weight. Measured in area. Mature in six
instars. Mature in seven instars.
instars of several species. The percentages are based on total food con-
sumption during the entire larval or nymphal stage. The great mass of
food is eaten during the last two or three instars. Bombyx and Proto-
parce, both lepidopterous leaf-feeders, eat about 97% of their total
intake during the last two instars and about 99% during the last three
i nstars.
The ideal utilization study will, of course, encompass the entire life
cycle. The efficiency of food utilization by the early instars is certainly
of interest. The nutritional adequacy of a food can be judged only by its
ability to support the growth of successive generations (Gordon, 1959).
However, studies which include only one or more of the later instars are
often useful. For instance, from an ecological point of view the rates of
digestion and conversion during the last two or three instars might
serve as a reasonably precise estimate of digestion and conversion
during the whole larval stage.
V . DIGESTIO
ANN D CONVERSIO
ONF F R E S HA N D D R Y
MATTER
A . LIMITATIONS OF THE DATA

Tables IV to VIII summarize much of the available informatioii on
the approximate digestibility and the efficiency of conversion of dry
matter and fresh matter to body substance by insects. The data in these
tables should be compared with discretion. First, apart from questions
of accuracy, data may not be strictly comparable for various reasons
which will be discussed below. Second, the data were obtained by
methods of varying accuracy. I have, however, omitted from the tables
data which appear to be grossly inaccurate or were not accompanied by
a statement of methods. I have also omitted data which are in no way
comparable with the main body of available information. For example,
Schwerdtfeger’s (1930) data are practically useless for comparative pur-
poses since he took only the fresh weight of the food, only the dry weight
of the feces, and used the increase in the length of his caterpillars as the
only measure of growth. McCay (1938) grew successive groups of
roaches on the same batch of diet until all of it had been consumed and
nothing but feces remained. He calculated the E.C.I. from the total
weight of roaches produced and the initial weight of the diet. The signi-
ficance of this figure is doubtful because of the high probability that the
roaches ate feces. Thus McCay’s (1938) E.C.I. probably falls somewhere
between a normal efficiency on a nutritious intake and a very low effi-
ciency on an intake consisting mostly of feces. The further digestion of
the feces adds additional complications.
Dry weight and fresh weight indices are not comparable, as can be
seen by examining corresponding indices derived from the same set of
C O N S U M P T I O N AND U T I L I Z A T I O N O F F O O D 25 1
insects in the same experiment. Corresponding dry weight (Table IV)
and fresh weight indices (Table V) are available for Bombyx, Malaco-
soma, Phalera, Aglais, Pieris, Dermestes, Tribolium and Anagasta.
Corresponding fresh and dry weight indices may differ to varying
degrees, the difference depending upon the water content of the food
and the insect and the % utilization of water by the insect. Fresh weight
E.C.D.’s for Dermestes, Anagasta and Tribolium are two to three times
higher than corresponding dry weight E.C.D.3. This probably reflects
a high rate of retention of the small amount of water taken in with the
food, and the storage of metabolic water obtained by the oxidation of
carbohydrates.
The data in Tables IV and V were taken over different lengths of time
and during different portions of the life cycle. It should be kept in mind
that the efficiency of utilization is likely to differ from stage to stage,
from instar to instar or even within an instar. Thus measures made over
a portion of an instar or stage are not necessarily representative of that
entire instar or stage (see Section VE). The data of Brennikre et al.
(1949), Teissier (1931) and Wolcott (1924) were further complicated by
their use of groups of insects of mixed ages.
Some of the efficiencies of conversion were calculated with gross
weight gain while others were calculated with net weight gain (see foot-
notes of Table IV). An efficiency calculated from gross gain (includes
weight lost during molts) will, of course, be higher than a corresponding
efficiencycalculated from net gain. For example, all of the efficiencies
of conversion of ingested food (E.C.I.) available for piercing-sucking
predators or blood-feeders are based on net gain (Table V). In each case
gain was determined by subtracting the weight of an individual before
feeding from its weight after feeding and molting to the following instar.
The efficiency of utilization will vary with temperature, relative
humidity and other physical factors. Teissier’s (1931) efficiencies of con-
version of digested food (E.C.D.) represent Tenebrio larvae kept at
temperatures well below the optimum, and eating either wheat flour or
powdered milk with suboptimum moisture contents, 12-5 and 23%,
respectively. These conditions were reflected in an extremely slow growth
rate and are probably also reflected in the E.C.D. (Table IV).
An insect may utilize a food which it does not normally eat far less
efficiently than its natural food. This has been demonstrated with
Prodenia (So0 Hoo and Fraenkel, 1966) and Protoparce (Waldbauer,
1964) (Table IV). Edwards (1964) found an extraordinarily low E.C.I.
of carrot root by the larvae of Hepia/us (Table V). Carne (1966) sug-
gested that carrots may not be one of the natural foods of Hepialus.
t
4
TABLE 1V VI
h,
The utilization of dry matter by insects
Temp. (C) Sex and Instar

Insect and authority and r.h. Food or Stage A.D. E.C.I. E.C.D.
Leaf Feeders
Locusta migratoria 32" Grass 3 Days early in instar V
(Orthoptera: Acrididae) 55% 39
Dadd (1 960)
Melanoplus bilituratus 30" Renown wheat Whole nymphal stage 32
(Orthoptera : Acrididae) - Triticuni nestivutn
Smith (1959) Agropyron smithii
western wheat grass As above 32
Ajax oats
Acena satica As above 32
Schistocerca gregaria 30' Scarole (escarole) Instar IV (7 days)
(Orthoptera : Acrididae) - Cichorirrni endiuia ? gregarious phase
Brennihre et al. (1949) (Compositae) isolated 55
gregarious phase
in groups 54
solitary phase 61
As above 30" As above Adult 6 (24 hr) 33
Chauvin (1946) - Adult 9 (24 hr) 33
A s above 32' 3 Days early
Dadd (1960) 55% Bran in instar V 40
Grass As above 39
As above 28-35" Mixed grasses, mostly Poa sp. Adult 6
Norris (1961) - and Loliiirii sp. 1st to 10th day
crowded 42"
isolated 3 6"
1 I th to 20th day
crowded 41"
isolated 38"
Gryilus domesticus 30' Scarole (escarole) Adult 8 (24 hr) 42
(Orthoptera: Gryllidae) - Cichoriunr endivia ?
Chauvin (1946) (Compositae) Adult p (24 hr) 37
Cararisius morosus 22" Hedera helix Instar IV 55"
(Orthoptera: Phasmidae) - (Araliaceae) (30 days)
Brenniitre et al. (1949)
As above 20" Hedera helix Whole nymphal
Lafon (1951) - (Araliaceae) stage+?) 51
Phaneropterafalcata 30" Lettuce
(Orthoptera : Tettigoniidae) - Lactuca satica Adult d (24 hr) 41
Chauvin (1946) (Compositae) Adult F (24 hr) 41
Hyphantria cunea - Acer negundo Whole larval stage 22
(Lepidoptera : Arctiidae) - (Aceraceae) C
Balogh and Gere (1953)b i
As above - Malus punrila Whole larval stage 31" - - rN
-
2
Nagy (1953) (Rosaceae)
Bontbyx mori 22" Morus alba Whole larval stage 37" 23" 62c
(Lepidoptera : Bom bycidae) - Goshoerami 0
Hiratsuka (1920) variety (Moraceae) z
As above - Morris alba Instar IV-V 39 19"" 48" 0
Shyamala et a / . (1960) - n
As above 27" Morus alba Instar IV 44 21 47
So0 Hoo and Fraenkel(l966) - 0
0
Malacosoma neustria - Salix uiminalis Part of (24 hr?) last instar 34 14" 41 tl
(Lepidoptera: Lasiocampidae) - (Salicaceae)
Evans (1939b)b
Lynrantria monacha 20-22" Fagirs sp. Instar IV 29
TABLE
Iv-continued
Temp. ((3 Sex and Instar

(Lepidoptera: Lymantriidae) ca 75% (Fagaceae) Instars V and VI 21 - -

Sattler (1939)b Alnus sp. Instar IV 35 - -
(Betulaceae)
Picea sp. Instars V and VI 25 - -
(Pinaceae)
Agrotis orthogonia - Sprouts of Thatcher wheat
(Lepidoptera: Noctuidae) - (Triticum aestivum) Instar Iv-VI 5oC 28" 56"
Kasting and McGinnis Sprouts of golden ball wheat
(1 959) (Triticum durum) Instar IV-VI 55" 26c 46c
Mamestra brassicae 25" Beet, Beta sp. Early instar V to pupa 49 14'." 29"
(Lepidoptera: Noctuidae) (Chenopodiaceae)
Hirano and Noguchi - sweet potato
(1963) Ipomoea batatas As above 24 3%e 14'
(Convolvulaceae)
Prodenia eridania 27" Morus alba Instar V 50 21 42
(Lepidoptera: Noctuidae) - Phaseolus lunatus
So0 Hoo and Fraenkel (Leguminosae)
(1966) Mean of data on
(For data of So0 Hoo and simple leaves As above 66" 29" 43"
Fraenkel, plants belong to trifoliate leaves As above 66 33 49
last family mentioned if P. vulgaris As above 70 38 54
family is not otherwise Taraxacum oficinale
indicated) (Compositae) As above 60 31 52
Antirrhinum majus
(Scrophulariaceae) As above 65 10 16
Ipomoea batatas
(Convolvulaceae) As above 40 18 44
Nicandra physalodes"
(Solanaceae) As above 66 20 30
Solanum tuberosum As above 60 34 57
Lycopcrsicon esculentirni As above 64 22 35
Abutilon theoplirasti"
(Malvaceae) As above 50 14 29
Chenopodium album
(Chenopodiaceae) As above 44 23 51
Spinacia olcracea As above 60 26 42
Phytolacca americana
(Phytolaccaceae) As above 58 30 52
Viola ~ p . ~
(Violaceae) As above 53 10 19
Malus floribunda"
(Rosaceae) As above 36 11 29
Prunus serotinaa As above 47 8 18
UImus pumila
(Ulmaceae) As above 44 11 25
Acer saccharinuni
(Aceraceae) As above 46 12 26
As above Phaseolus vulgaris Instars V and V1 48 34" 69"
Crowell (1941)b London Hort. variety
(Leguminosae)
Phalera bucephala Carpinus betula Part of (24 hr?) last instar 35 - -
(Lepidoptera: Notodontidae) (Betulaceae)
Evans (1939b)b
As above Corylus avellana 24 hr? instar? 35 - -
Evans (1939a) (Betulaceae)
Aglais urticae Urtica dioica Part of (24 hr?) last instar 26 17c 63
(Lepidoptera: Nymphalidae) (Urticaceae)
Evans (1939b)b
Pieris brassicae Cabbage
TABLE
IV-continued h,
wl
o\
Temp. (0 Sex and Instar

(Lepidoptera: Pieridae) - Brassica oleracea Part of (24 hr?) last instar

Evans (1939b)b (Cruciferae)
young leaves 36 -
old leaves As above 39 -
Chilo suppressalis 28" Steam sterilized rice stems First 20 days of larval 57 13c*"
(Lepidoptera: Pyralidae) - stage, ca instar I-V
Hirano and Ishii (1962)
Protoparce sexta - Tobacco
(Lepidoptera : Sphingidae) - Nicotiana sp. Instar IV 50" 25"
Wolcott (1937)b (Solanaceae)
As above 26-30' Lycopersicon esculentuni Instar I V 49" 27"
Waldbauer (1 964) - (Solanaceae)
Mean of all data
Solanum tuberosum As above 56 36
(For data of Waldbauer, S. dulcamara As above 42 27
plants belong to last family Taraxacum oficinaled As above 39 21
mentioned if family is not (Compositae)
otherwise indicated) Arctirim minusd As above 41 20
Verbascuni thapsusa*
(Scrophulariaceae) As above 32 12
Non-Leaf Feeders
BIatta orientalis 20" Powdered milk, yeast and Last 2 nymphal instars 95 -
(Orthoptera: Blattidae) - cholesterol
Lafon (1951)
As above 25' Wheat flour Adult d (24 hr) 63 -
Chauvin (1 946) - Adult 0 (24 hr) 59 -
Gryllus domesticus 30" Wheat flour Adult d (24 hr) -
(Orthoptera : Gryllidae) - Adult 0 (24 hr) -
Chauvin (1946)
As above 30" Wheat flour Nymphs of 118-272 mg 1 9"
Brennitre et al. (1949) - (1 5 days)
Dermestes maculatus 25" Brewer's yeast, fructose and
(Coleoptera : Dermestidae) 70% cholesterol Instar I to pupa 19"
Fraenkel and Blewett 50% As above As above 16"
(1944)b 30% As above As above 11'
Tenebrio molitor
(Coleoptera : Tenebrionidae) 27" Wheat bran Larvae (4 days) -
Evans and Goodliffe (1939) -
As above 17" Wheat flour Larvae of mixed ages 1 8"
Teissier (1931)b - (18-45 days)
20" Powdered milk As above (14-1 6 days) 23"
Tribolium confusuni 25" White flour and yeast
(Coleoptera : Tenebrionidae) 70% Instar I to pupa 19"
(1944)b
Anagasta kuehniella 25" Whole wheat flour
[ =Ephestia] 70% Instar I to pupa 16"
(Lepidoptera: Phycitidae) 20% As above As above 11"
Fraenkel and Blewett > 0% As above As above 8"
(1944)b
A.D. = approximate digestibility PA),E.C.I. = efficiency of conversion of ingested food to body substance, E.C.D. = efficiency of conversion
of digested food to body substance.
The E.C.I. and E.C.D.are based on gross gain unless otherwise indicated. The A.D. and E.C.D. have not been corrected for uric acid content
of the feces.
Unless otherwise indicated experiments ran from the beginning to the end of the instar or stage mentioned or from the beginning of the first
to the end of the last instar mentioned.
8 Plants which do not or only rarely support growth. See text for discussion of author's methods or other comments. Calculated from
author's data. Plants not usually eateniaccepied only after maxillectomy. Based on net weight gain. Exuviae included in weight of feces.
V
TABLE h,
ch
00
The utilization of fresh weight of food by insects

Leaf Feeders
Carausius morosiis - Hedera helix Whole nymphal stage ( 0 ) -
(Orthoptera: Phasmidae) - (Araliaceae) Whole adult stage (0) -
Titschack (1924)b
Paropsis atomaria - Ei4calyptus blakelyi Instar I to prepupa 20"
(Coleoptera: Chrysomelidae) - young shoots
Carne (1 966) (Eucalyptaceae)
Bombyx mori 22" Morus alba Whole larval stage 25"
(Lepidoptera: Bombycidae) - Goshoerami variety
Hiratsuka (1920) (Moraceae)
Hepialus humuli 20" Slices of carrot root Instar I to pupa 2"
(Lepidoptera: Hepialidae) 98%
Edwards (1964)b
Malacosoma neustria - Salix uiminalis Part of (24 hr?) last instar 22'
(Lepidoptera: Lasiocampidae) - (Salicaceae)
Evans (1939b)b
Dendrolimuspini - Pinirs sp. Whole larval stage 13'
(Lepidoptera: Lasiocampidae) - (Pinaceae)
Lebedev and Savenkov
(1932)"
Phalera bucephala - Carpinus betula Part of (24 hr?) last instar 28"
(Lepidoptera: Notodontidae) - (Betulaceae)
Evans (1939b)b
As above - Corylus avellana 24 hr? instar? 34
Evans (1939a)" - (Betulaceae)
Aglais urticae Urtica dioica Part of (24 hr?) last instar 24 14
(Lepidoptera: Nymphalidae) (Urt icaceae)
Evans (1939b)"
Pieris brassicae Cabbage
(Lepidoptera: Pieridae) Brassica oferacea
Evans (1939bIb (Cruciferae)
Young leaves Part of (24 hr?) last instar 20 15c
Old leaves As above 24 16"
As above Cabbage
David and Gardiner Brassica oleracea Whole larval stage - 13"s"
(1962)b
Snterinthus populi Salix sp. First instar 64" 32c
(Lepidoptera: Sphingidae) (Salicaceae)
Evans (1939b)b
Non-leaf Feeders
Cryptotermes brevis Seasoned, air-dry wood Mixed stages
(Isoptera: Kalotermitidae) Asubo
Wolcott (1924)b Sideroxylon foetidissimrni 12 -
Pomarrosa
Eugenia jambos 41 -
Sitka spruce
Picea sitchensis 51 -
Flamboyan
Poinciana renia 58 -
Cimex lectularius Blood of hum& 1 Instar 11 (unfed) to 0
instar V (unfed) - - 0
(Hemiptera: Cimicidae) 3oc.e
t3
Johnson (1960)" AS above Blood of human 2 As above - 3oc.e -
As above Blood of mouse As above - 3oc.e -
As above Blood of fowl Instar 11(unfed) to
instar IV (unfed) - 3W" - N
TABLE
V-continued
E
30" Blood of man Instar I (unfed) to

92% instar V (unfed) -
20" Asabove Instar I (unfed) to
86% adult (unfed) -
Stalia major 13.5-1 9" Larvae of Carpopidus Jnstar I (unfed) to
(Hemiptera: Nabidae) and dimidiatus instar V (unfed) -
Fewkes (1960) 20-22" (Coleoptera: Nitidulidae) 0
(pooled cd
results of two
experiments) E
Rhodnius prolixus 26-27" Rabbit blood Instar I (unfed) to 9
(Hemiptera : Reduviidae) - instar V (unfed) - r
tl
Friend et al. (1965)b W
9
sP
Phonoctonus nigrofasciatus 27" Nymphs of Dysdercus fasciatits Instar I (unfed) to
(Hemiptera : Reduviidae) 75:4 (Hemiptera: Pyrrhocoridae) instar V (unfed) -
Evans (1 962)
Sitophilus granarius 25" Whole grains of wheat Instar I to adult 14e
(Coleoptera: Curculionidae) 70%
Richards (1947)
Dertnestes maculatus 25" Brewer's yeast, fructose and
(Coleoptera : Dermestidae) 70% cholesterol Instar I to pupa 38"
Fraenkel and Blewett 50% Asabove As above 35"
(1944)b 30% Asabove As above 36"
Tribolium confusutii 25" White flour and yeast
(Coleoptera : Tenebrionidae) 70% Instar I to pupa 39"
(1 944)b
c)
0
z
(A
C
3
%!
25" Whole wheat flour r!
Anagasta kuehniella
[ = E'hestia] 70% Instar I to pupa -
-
-
-
42' 2
30"
(Lepidoptera: Phycitidaz)
Fraenkel and Blewett
20%
iO%
As above
Asabove
As above
As above - - 22" ;
(1944)b t,
Tineola bisselliella 30" Wool dipped in brew of
'
C
(Lepidoptera: Tineidae) horse feces Instar I to adult ( 6 ) 39" 16".' 41C*e 2
Titschack (1925)" 30" As above As above ( 0 ) 41" 38's"
20" As above As above (d) 45' 18C.e w.e
N
20" As above As above ( 0 ) 45" 22C.' 49c.e >
2
0
See Table IV for footnotes. z
C
7
?I
0
0
0
Beck (1956) pointed out some of the difficulties which may be involved
in the use of excised plant tissues in nutritional studies. Excised tissue
may change rapidly because of biochemical degradation, changes in
water and hydrogen ion relationships and microbial attack.
Some of the data quoted in the tables were obtained by methods
which may have led to relatively minor inaccuracies. Fraenkel and
Blewett (1944) were unable to separate the silk spun by Anagasta
kuehniella larvae from the mixture of uneaten food and feces. This
makes their figure for food digested somewhat too low and thus inflates
the E.C.D. Titschack (1924), Crowell (1941), Husain et al. (1946) and
David and Gardiner (1962) calculated the weight of food eaten from the
area eaten. This was probably reasonably accurate if there is little varia-
tion in the relationship of area to weight. Sattler (1939) used woody
twigs with attached leaves both as food and aliquots for dry weight
determinations. He selected twigs which appeared to have woody por-
tions of the same length and thickness, and trimmed enough leaf material
from each so that all had the same fresh weight. Variations in the pro-
portion of woody material may have made his dry weight determinations
somewhat inaccurate. Lebedev and Savenkov (1932) determined the
number of pine needles which had been eaten and calculated their
weight from the average weight of a needle which had been previously
determined from an aliquot of needles taken from the same twig. Davey
(1954) used the % dry matter of the grass left uneaten after 24 hr to
estimate the dry weight of the grass initially provided. This probably
inflated his figure for the dry weight of grass provided, thus deflating
the A.D. Balogh and Gere (1953) took the air-dry weight of food and
feces. Their figures for consumption and excretion are, therefore, prob-
ably somewhat higher than those of workers who oven-dried this
material. A.D.’s determined on a fresh weight basis will be somewhat
low if there was a loss of moisture from the feces. Data for small species
or early instars may be less accurate than data for larger forms because
of the small quantities involved.
Hiratsuka (1920) states that the “average digestibility” of dry matter
during the whole larval stage of Bornbyx is 40.9%. An examination of
his data reveals that this figure was indeed arrived at by taking the mean
of the A.D.’s for the five instars. This average figure is not the same as
the % digestibility of the total amount of food consumed during the
larval stage. A.D. is at its highest in the first instar, but the food intake
of this instar is only 0.08% of the total. The food intake of the 5th
instar is 85% of the total, but A.D. is at its lowest in this instar (Table
VII). Thus the mean is biased by a high A.D. at a time of low intake.
Calculating from the total weights of food eaten and feces passed during
the entire larval stage gives a considerably lower A.D. of 37%. Several
authors calculated similar “average values”. In each case I have cal-
culated the actual overall value for presentation in this paper.
B . COMPARISON OF SPECIES
A glance at Tables IV and V shows that approximate digestibility and
efficiency of conversion differ widely from species to species. On the
whole it is probably a safe assumption that these differences are largely
the result of differences inherent in the normal insect-food relationship.
However, as pointed out above, comparison is complicated by varia-
tions in accuracy, experimental design, environmental conditions and
other factors.
The data of Table IV suggest that leaf-feeding Lepidoptera may in
general utilize food for growth more efficiently than the nymphs of leaf-
feeding Orthoptera. Unfortunately, all three indices are available for
only eight of the Lepidoptera and three of the Orthoptera. Considering
only these species and omitting the nutritionally inadequate foods, it
can be seen that the Lepidoptera on the whole convert ingested food to
body substance (E.C.I.) more than twice as efficiently as the Orthoptera.
The difference must be attributed largely to the greater efficiency of
conversion of digested food (E.C.D.) by the Lepidoptera, since approxi-
mate digestibility does not differ much in the two groups. Of the Lepi-
doptera under consideration only Mamestra and Prodenia on a few of
its foods show low E.C.D.’s.
So0 Hoo and Fraenkel (1966) compared the utilization of mulberry
(Morus) leaves by Bombyx and Prodenia under nearly identical experi-
mental conditions. They used the penultimate larval instar of each
species, 4th instar Bombyx and 5th instar Prodenia. The E.C.I.’s were
the same. However, the higher A.D. in Prodenia compensated for its
lower E.C.D. (Table IV). The data of So0 Hoo and Fraenkel(l966) and
Waldbauer (1964) for Protoparce permit similar but somewhat less pre-
cise comparisons (Table IV). On tomato (Lycopersicon) Prodenia has a
somewhat lower E.C.I. than Protoparce. However, the considerably
higher A.D. in Prodenia again compensates for its far lower E.C.D.
A similar situation pertained when potato (Solanum tuberosum) leaves
were fed. On Taraxacum leaves Prodenia had the higher E.C.I., but this
was clearly the result of a higher A.D. Taraxacum is not a natural food
for Protoparce and usually is accepted only by maxillectomized larvae.
These observations are of considerable interest. Both Bombyx and
Protoparce are oligophagous. Bombyx will normally eat only mulberry
or a few other species of Moraceae. Protoparce in nature eats tobacco or

tomato, but will accept other Solanaceae in the laboratory. Prodenia, on
the other hand, is polyphagous and readily accepts a wide variety of
plants. One would expect Bombyx and Protoparce to be specifically
adapted to digest and convert their limited diets. On the other hand, one
would expect Prodenia to be more generalized, adapted for the reason-
ably efficient utilization of a variety of foods. Thus the comparatively
high E.C.D.’s in Bombyx and Protoparce suggest specific adaptations
to the nutrient balances offered by their natural foods. The low E.C.D.
in Prodenia suggests a less precise correspondence between its require-
ments and the nutrient balance of its diet. Perhaps the high A.D. in
Prodenia is an adaptation which compensates for a decrease in the effi-
ciency of conversion resulting from moderate nutritional imbalances.
C . C O M P A R I S O N O F FOODS
Several comparisons of the utilization of different foods by the same

species are available. I refer the reader to the data for Melanoplus,
Gryllus, Lymantria, Mamestra, Prodenia (So0 Hoo and Fraenkel, 1966)
and Protoparce (Waldbauer, 1964) in Table IV and Cryptotermes and
Cimex in Table V.
It is clear that at least among the leaf-feeding species digestibility and
efficiency of conversion vary widely with the species of food plant.
Maxillectomized Protoparce larvae will feed on some normally rejected,
non-solanaceous plants (Waldbauer and Fraenkel, 1961). Some non-
solanaceous plants were efficiently utilized while others did not support
normal growth although they were eaten at the same or higher rates
than plants which supported good growth (Waldbauer, 1964) (Tables I1
and IV). So0 Hoo and Fraenkel(l966) obtained similar results with the
intact polyphagous larvae of Prodenia (Table IV). These findings are
contrary to Fraenkel’s (1959) earlier contention that the leaves of all
plants are of similar nutritional value. They do not, however, conflict
in any way with Fraenkel’s (1959) theory of host plant selection, that
secondary plant substances serve as “ token feeding stimuli” (sign
stimuli).
Gryllus adults digested wheat flour at a much higher rate than
escarole leaves (Chauvin, 1946) (Table 1V). This is not surprising since
the crude fiber content as a percentage of dry matter is lower in flour
than escarole. Wolcott (1924) found that the approximate digestibility
of various woods by Cryptotermes ranges from 12 to 58% (Table V).
Cimex converts the blood of humans, mice or fowl with equal efficiency.
In each case the E.C.T. was 30% (Johnson, 1960) (Table V). Unfortu-
nately the other indices are not available for any of these insects.
It is also clear that leaves of the same species of plant may vary in
nutritional value. So0 Hoo and Fraenkel (1966) measured the utiliza-
tion of Phaseolus lunatus leaves by Prodenia eleven times. The means of
these measurements are given in Table IV. In these tests, made at
different times of the year, A.D. ranged from 56 to 72%, E.C.I. from
19 to 38% and E.C.D. from 33 to 52%. The utilization of tomato leaves
by Protoparce seems to be less variable. The means of data obtained
from four experiments conducted in July are given in Table IV. A.D.
ranged from 43 to 52%, E.C.T. from 24 to 30% and E.C.D. from 55 to
59%. Waldbauer (1964) thought that some of the variation may have
been due to differences in the water content of the field-grown leaves.
In both Prodenia and Protoparce the variation in the utilization of
different species of plants is greater than the variation in the utilization
of different samples of the same plant species. Evans (1939b) found that
for Pieris old cabbage leaves are more digestible than young leaves, and
that both are converted to body substance with almost equal efficiency
(Tables IV and V).
D . EFFECTS O F ENVIRONMENTAL FACTORS
A large body of literature shows that mortality, growth, fecundity and
other of the vital statistics of insects vary widely with environmental
factors. However, almost nothing is known of the effects of the environ-
ment on the rate of consumption, digestibility, or the efficiency of food
conversion. Existing data suggest that relatively small changes in the
physical or biotic environment may have significant effects. The dis-
covery and elucidation of these effects should prove to be an interesting
and fertile field of research.
The amount of food consumed and the efficiency of its utilization
may vary with the degree of crowding. Crowded Schistocerca gregaria
nymphs consumed more food than nymphs reared in groups of three
(Davey, 1954). Norris (1961) found that A.D. is higher in crowded
Schistocerca adults than in isolated adults. However, Brennittre et a/.
(1949) found little difference between the A.D.’s, E.C.T.’s and E.C.D.’s
of 4th instar gregarious phase Schistocerca raised in crowds or in isola-
tion (Table IV). Hyphantria cunea larvae reared individually or in small
groups grew somewhat more slowly than larvae reared in large groups
(Gere, 1956). Long (1953) found that crowded Plusia gamma completed
the larval stage more quickly than isolated individuals, although isolated
individuals appeared to gain more weight. Furthermore, the relative
effect of crowding was the greatest on the nutritionally best plants.

Crowding led to the largest proportional increase in the rate of develop-
ment on those plants which supported the most rapid growth of isolated
larvae.
The effect of temperature on the efficiency of Tineolu and Hyphuntria
is discussed below (Section V E ~ )The
. amount of food consumed by
Bombyx larvae varies widely with temperature. Taking the amount
consumed in a given time at 22" as loo%, consumption fell to 46% at
14' and rose to 203% at 32-33'. The A.D. did not vary greatly from 18
to 32', but it was 44% lower at 14" (Legay, 1957). Sattler (1939) found
that the A.D. of Lymuntria fell with increasing temperatures above 14',
but that the magnitude of the fall varied with the food plant and the
instar (Table VI).
TABLE
VI
The effect of temperature on the approximate digestibility (%) by Lymanfria
monacha of the leaves of Alnus incana, Fagus sylvaticus, and Picea excelsas
4th instar 5th and 6th instars

Temperature "C Alnus Fagus Fagirs Picea
12-14 60 32 28 44
20-22 36 29 21 25
24-26 30 30 22 18
a From Sattler (1939).
Little is known concerning the effecton food intake and utilization of

relative humidity and other factors which influence water loss from the
body. Many insects are known to drink (Barton-Browne, 1964), but
others, particularly stored-product pests, and perhaps many leaf-feeders,
probably get all or most of their water in the food. Water loss from the
body would presumably be compensated for by increasing food intake
-either for its water content or as a source of metabolic water. In either
case increased intake would probably lead to a decrease in the efficiency
of utilization.
Fraenkel and Blewett (1944) found that Anugasta, Dermestes and
Tribolium, pests of stored dry products, convert less of the digested food
to body substance as the relative humidity of the air and the water con-
tent of the food are decreased (Tables IV and V). A larger amount of
food was digested at the lower relative humidities, but much of it was
used as a source of metabolic water. The decrease in efficiency probably
involved not only the loss of the carbohydrates metabolized for water,
but also a corresponding waste of digested nitrogen.
Larvae of Diataraxia oleracea grew normally and did not drink free
water when they were fed turgid cabbage leaves which contained about
85% water. However, when fed wilted cabbage leaves (70% water or
more) they became dehydrated and drank large quantities of water
(Mellanby and French, 1958). Legay (1957) found that Bombyx larvae
fed mulberry leaves containing at least 70% water retained much of the
ingested water. About 30% was passed out with the feces, 10% was lost
as vapor and 60% was stored. The data of Sharada and Bhat (1957)
suggest that Bonibyx larvae may digest and convert both dry matter
and nitrogen more efficiently at 1000/,r.h. than at a lower but unspecified
ambient r.h. Evans (1939b) stated that lepidopterous larvae which ate
leaves with a high water content ( > 60-90%) absorbed relatively little
of the water ingested with the food (20-30%), while larvae which ate
leaves with a low water content ( < 60- > 50%) absorbed much more
water (6040%).
Legay (1957) commented that a study of utilization based entirely on
dry weights ignores the important question of water balance. However,
we have little quantitative information on the intake and utilization of
water by insects. The data will be difficult to obtain. Perhaps the most
difficult problem will be the accurate determination of the fresh weight
of feces.
E . V A R I A T I O N S W I T H A G E A N D SEX
1. Variation with age

Approximate digestibility and efficiency of conversion apparently do
not remain constant, but rather vary with age during the growth period
of an insect. Table VII summarizes the data which permit comparison
of A.D. from instar to instar. In each case A.D. declines with age.
Evans’ (1939b) data for Phalera bucephala do not show this tendency as
clearly as do the rest, probably because he did not measure A.D. over
the entire instar, but sampled it for 24 hr periods at intervals of from
one to nine days. This could yield figures not representative of entire
instars since, as will be shown below, A.D. varies within instars. Cal-
culations made from the data of Kasting and McGinnis (1959) show the
A.D. for Agrotis remaining more or less constant in the last three
instars. They stated that their data for the first three instars are not
reliable. This does appear to be the case. In two instances calculations
from their data yield impossible percentages of digested food retained
W
TABLE
The A.D. from instar to instar
I
Instar
Insect, food, and authority I I1 III IV V
Bombyx mori
Leaves of Morus alba 47 44 38 38 37
Hiratsuka (1920)
Plialera bucephala
Leaves of Carpinits hefula
Dry weightR 24 18 22 24 19
Fresh weight” 70 43 42 46 45
Evans ( 1939b)
Scliltocerca gregariu
Mixed grasses 78 52 45 34 35
Davey ( 1954)”
S. gregaria
Leaves of Ficits sp. 48 42 45 38 31
Husain et a / . (1946)”
All figures are OR a dry weight basis unless otherwise indicated.

a Means of A.D.’s measured over 24 hr at intervals of several days, probably not repre-
sentative of the A.D. for the entire instar.
See text for discussion of author’s methods or other comments.
as body substance, 133% for 1st instar larvae feeding on Golden Ball
wheat and 122% for 2nd instars feeding on Thatcher wheat.
Other workers who did not give data for individual instars also found
that A.D. declines during the larval or nymphal stage. Dry weight A.D.’s
for sixteen of the twenty days of the larval stage of Hyphantria can be
calculated from Nagy’s (1953) data. The A.D. of the leaves of Malus
pumila is 49 on the first day, fluctuates about a mean of 34 for the next
fourteen days, and falls to 12 during the last five days. However, calcu-
lations from the data of Balogh and Gere (1953) show dry weight
A.D.’s of Acer negundo leaves for Hyphantria larvae of 39 for the first
fourteen days, 19 for the next ten days, and 23 for the last eight days.
Smith (1959) found that the dry weight A.D.’s of Agropyron smithii
(western wheat grass), Triticum aestiuurn (wheat) or Auena satiua (oats)
for nymphs of Melanoplus bilituratus decreased during the first twenty
days, tended to level off during the next ten days, and decreased again
during the last ten days of the nymphal stage. The initial A.D.’s of the
grass, wheat and oats were 81, 65 and 54, respectively. They fell to
between 20 and 30. Sylven (1947) found that the larvae of Phyfometra
gamma digest less of soya bean leaves with increasing age. He measured
consumption in square millimeters and feces in dry weight, but his find-
ings are probably valid for this comparison. Calculations from the data
of Schwerdtfeger (1930) show that the digestibility of pine needles for
the larvae of Bupalus piniarius decreased gradually as they grew from a
length of 2.8 to 29-3mm during the larval stage of seventy-two days.
He measured consumption in fresh weight and feces in dry weight, but
his figures are also probably valid for this comparison.
The reasons for the decline of A.D.are not entirely clear. One would
expect digestive efficiency to decline with growth since an animal which
doubles its weight and volume will increase the surface area of the
digestive tract by a factor of only 1.8 (Gordon, 1959). With chewing
insects it could be argued that small individuals chew off smaller pieces
of food and thus present a greater surface area for digestion. Small leaf-
feeders might also ingest a greater proportion of easily digested broken
cells (Biedermann, 1919). Evans (1939b) pointed out that the composi-
tion of the food selected by leaf-feeders changes as they grow older.
First instar larvae eat from between the small veins of the leaf. Some-
what older larvae also eat the small veins, while larger larvae eat almost
the whole leaf. Thus it is likely that the older larvae ingest a larger pro-
portion of indigestible crude fiber.
A.D. also varies within instars. The dry weight A.D. for Bombyx
varies as follows in the 5th instar: days 1 and 2,40%; day 3,47%; day
4,42%; day 5,39%; and on days 6 and 7 27% (Hiratsuka, 1920). Cal-
culations from the data of Evans (1939b) show that the fresh weight
A.D. of Salix leaves for a group of Smerinthus populi larvae varied as
follows from the first to the tenth day of the 1st instar: 92,74,61,56,56,
56, 59, 52, 50 and 53%. Many workers have observed that leaf-feeding
insects pass more and wetter and greener feces near the end of an instar,
especially the last instar. This is probably a sign of a decrease in A.D.
Table VIII is a summary comparison of the variation of the efficiency
of conversion of ingested food (E.C.I.) from instar to instar. With the
exception of Fewkes’ (1960) data for Stalia major there is an obvious
tendency for the E.C.I. to decrease with age. The E.C.I. of Stalia
nymphs, however, increased with age, but Fewkes (1960) thought that
this may have been because all instars were kept in cages of the same
size. The ratio of search area to the size of the predator was, therefore,
higher for the early than for the lateinstars. Thus the early instars
probably expended more energy to obtain each meal.
According to Edwards (1964) the fresh weight E.C.J. of Hepialus
hzrmzrli rises during the early instars, reaches more or less of a plateau
270 G . 1'. W A L D B A U E R
TABLE VILI
The E.C.I. from instar to instar
Instar
Insect, food and authority I rr [TI IV v VI
_- ~ -
Agrotis orthogonia*
Triticum aestiuunz sprouts - 35" 33" 27" 28" 17"
Triticum durum sprouts - 37" 26" 21" 22" 16"
McGinnis and Kasting (1959)
Bombyx morid
Morus alba leaves 31 27 25 25 23 -
Hiratsuka (1920)
Ciniex 1ectulariu.P-
Blood of man 41" 33" 31" 32" 24' -
Blood of mouse - 33" 34" 28" - -
Johnson (1960)
Phonoctonus nigrofasciatuP.
Dysdercus fasciatus nymphs 53" 54 43 42 33 -
Evans (1 962)
Rliodnius prolixusc*
Blood of rabbit 22 24 24 19 - -
Friend e f al. (1965)
Stalia majorc*
Carpophilus dimidiatus larvae 43 41 51 52 54 -
Fewkes (1960)b
~~ ~~ ~~ ~~
Indices are based on gross weight gain unless otherwise indicated.

* Calculated from author's data. See text for discussion of author's methods or other
comments. Fresh weight basis. Dry weight basis. Based on net weight gain.
and then drops to its lowest in the 12th instar. The significance of his
data is doubtful since slices of carrot root are apparently an unnatural
and a very poor food for this insect. The E.C.I.3 are exceptionally low,
ranging from 0.58 to 4-13. Calculations from the data of Evans (1939b)
suggest that the fresh weight E.C.I. of Phuleru increased from the 1st to
the 4th instar and decreased in the 5th. However, as pointed out above,
his data are probably not representative of entire instars since they are
based on eleven 24 hr measurements scattered over a larval stage of
forty-five days. Recalculation of the data of Carne (1966) shows that
the fresh weight E.C.I.'s of Puropsis utomuriu feeding on shoots of
Eucalyptus blukelyi are 18, 34, 30 and 18 from the 1st to the 4th instar.
There is little information on variation of the E.C.D. from instar to
instar. Hiratsuka (1920) found the E.C.D. of Bombyx to be 66, 61, 66,
65 and 61 from the 1st to the 5th instar, respectively. Calculations from
Evans' (1939b) data show the fresh weight E.C.D. of Phalera rising
from the 1st to the 4th instar and dropping sharply in the last: 21, 48,
68, 72 and 43, respectively. His data are, however, of doubtful signifi-
cance as stated above. Calculations from Smith's (1959) data show the
E.C.D. rising early in the nymphal stage and falling near the end of the
stage.
The decline of the E.C.I. with age is at least partly the result of the
concomitant decline of the A.D. Comparison of Hiratsuka's (1920)
indices for the five instars of Bombyx shows that the E.C.I.'s and the
A.D.'s decline together and at about the same rate. The E.C.D.'s, how-
ever, do not show a tendency to decline with age. Thus, at least with
Bombyx, the decline of the E.C.I. seems to be due to the decline in
digestibility and not to a decline in the efficiency of conversion of
digested food.
The E.C.I. can also vary within an instar. Carne (1966) reported a
rise and fall of the fresh weight E.C.I. from the beginning to the end of
the 4th and last instar of Paroysis, 19, 22, 18 and 4 on the 13th, 14th,
15th and 18th days of the larval stage, respectively. He suggested that
decreases in the E.C.I. are associated with energy-consuming physio-
logical activities associated with recent molts and the approach of
maturity. Calculations of Evans' (1939b) data for the 1st instar larvae of
Smerinthus populi show the E.C.I. varying from the 1st to the 10th day
of the instar as follows: 17, 28, 39, 39, 37, 36, 32, 36, 32 and 28.
2. Variation with sex
The sexes may differ in the efficiency with which they utilize food, but
whether or not sexual differences are of general occurrence is not clear.
Adult Gryllus males apparently digest more of either escarole leaves or
wheat flour than the females. The A.D. of wheat flour also seems to be
greater in adult males of Blattella (Table IV). In Tineola raised from
hatching to the adult stage at 30" approximate digestibility and net
efficiency of conversion differ only slightly from male to female. At 20"
both sexes digest and convert more efficiently, but although A.D. is the
same in both sexes, the E.C.I. of the female is higher because of an
increase in E.C.D. (Table V). Macko and Jasic (1959) measured A.D.
in the last two larval instars of Hyphantria reared at different tempera-
tures and feeding on the leaves of different species of trees. Approximate
digestibility of the more digestible leaves was consistently higher in the
female than the male. However, the sexes differed far less in their ability
to digest the less digestible leaves. The A.D.'s of both sexes varied with
temperature, but the % difference between corresponding A.D.'s of
212 G . P. W A L D B A U E R
female and male remained almost constant. In other species efficiency

does not seem to differ with sex. Chauvin (1946) found A.D. to be the
same in adult males and females of Schistocerca and Phaneropteru
.falcata (Table IV). Davey (1954) reported that total food intake during
the nymphal stage is greater in the female than the male of Schistocerca.
In the 5th instar the A.D., E.C.I. and E.C.D. of male and female Bombyx
larvae are similar, although the females seem to convert a little more
efficiently than the males (Hiratsuka, 1920). Legay’s (1957) findings with
Bombyx are similar.
V I . U T I L I Z A T I OONF D I E T A R YC O N S T I T U E N T S
The literature offers but few data on the utilization by insects of thc
nutrient constituents of either natural foods or chemically defined diets.
House (1959, 1962 and 1965a) pointed out that quantitative work with
defined diets has generally consisted of determining the amount of a
nutrient required per weight or volume of diet. Minimal or optimum
requirements expressed in these terms say nothing about the absolute
quantities required, but define only the relationships between the require-
ments for particular nutrients (Sang, 1956, 1959). This sort of informa-
tion facilitates the formulation of diets, but conveys a minimum of
biological meaning pertinent to questions of metabolism, ecological
relationships or the adaptation of insects to their natural foods. House
(1962) felt that absolute requirements have been neglected because their
determination requires the laborious task of measuring food intake.
The coeficient of apparent digestibility for the constituent X of the
food is usually expressed as a percentage and is calculated as:
amount of X in food ingested - amount of X in feces
C.A.D.(X) =
amount of X in food ingested
The eficiency with which the digestible portion of the constituent X is
converted to body substance is calculated as :
amount of X in body
E.C.D.(X) = x 100
- amount of X in feces
The eflciency of coizversiori of ingested constituent X to body substance
is calculated as:
amount of X in body
E.C.I.(X) =
Measurements of the utilization of nitrogen (N) by insects are compli-
cated by the presence of the urine in their feces. The feces, thus, contain
all of the absorbed N which was metabolized and excreted as waste.
Thus, if the amount of urine N is not determined and subtracted from
total fecal N, calculation by difference (N in food eaten less N in feces)
yields total N retained in the body rather than the amount of N digested.
The C.A.D. (N) and the E.C.D. (N) are, therefore, practically meaning-
less if they are calculated from data which have not been corrected for
urine N. The uncorrected C.A.D. (N) and the E.C.I. (N) should, of
course, be identical since ingested N less fecal and urine N should be
the same as the amount of N retained in the body. The uncorrected
E.C.D. (N) should always be 100% for the same reason. Both un-
corrected calculations are meaningless truisms.
The last column of Table JX shows that most of the E.C.D.’s (N)
actually deviate considerably from the expected 100%. It can be reason-
ably assumed that these deviations are due to the incomplete recovery of
N during analysis. The uncorrected E.C.D. (N) thus serves as a measure
of the accuracy of the N determination. Balogh and Gere (1953), how-
ever, ascribed an apparent surplus in N retention to the fixation of
atmospheric N.
An approximate correction for urine N can be made for only three of
the species listed in Table IX. Evans and Goodliffe (1939) and Hiratsuka
(1920) gave the uric acid content of the feces and Evans (1939a) gave
both the uric acid and ammonia content. Other nitrogenous wastes are
not accounted for, but with the insects in question correction for uric
acid alone probably closely approximates correction for total urine N.
Available analyses show that, except in blowfly larvae and some
aquatic larvae, uric acid generally accounts for 80% or more of the
excreted N (Stobbart and Shaw, 1964). The resulting difference between
the corrected and uncorrected C.A.D.3 (N) varies. With Bonibyx the
corrected C.A.D. (N) is about 7”/, higher, with Phalera about 15% and
with Tenebrio about 38%. Calculations from the data of Hiratsuka
(1920) give a corrected E.C.D. (N) of 92% for Bornbyx as opposed to
the uncorrected value of 100.4% (Table IX).
It should be noted that even if all of the urine N is accounted for the
apparent rather than the true digestibility of N is obtained because of
the fecal metabolic N (is. peritrophic membrane, see Section 11B2). The
accuracy of E.C.J. (N) is not, of course, affected by the presence of urine
or other metabolic N in the feces.
Whether an efficiency of conversion is based on the gross retention or
the net retention of N may make a considerable difference. The former
TABUIX
r4
The utilization of nitrogen 4
P
C.A.D. % E.C.D. E.C.D.
(Total N) (Total N) (Total N)
C.A.D. % corrected for E.C.I. corrected for not corrected
Insect, food and authority (Protein N) uric acid (Total N) uric acid for uric acid
Agrotis orthogonia
Thatcher wheat sprouts 1158*c
Golden ball wheat sprouts 95".
Kasting and McGinnis (1959)
Bombyx mori
Morus alba leaves 78" 63" 608 92".
Hiratsuka (1920)
Chilo suppressalis
Autoclaved rice stems 70 - 62"
Hyphantria cunea
Acer ncgicndo leaves - 44" 1 14"
u0
Balogh and Gere (1953)
Mamestra brassicac crn
P
Beet leaves - 31" 97" w

Sweet potato leaves - 12" 94"
Hirano and Noguchi (1963)
Phalera bucephala
Corylus avellana leaves 56
Evans (1939a)
Tenebrio molitor
Wheat bran 54 -
Evans and Goodliffe (1939)
The experiments in which these data were obtained are summarized in Table IV. E.C.I.(N) and E.C.D.(N) are based on net retention
of N unless otherwise noted.
a Calculated from author's data. Corrected for ammonia in feces in addition to uric acid. Based on gross retention of N; thus N
lost in the exuviae at the molt is included.
includes the N content of the exuviae and products such as silk while the
latter does not. The N content of the exuviae of Bombyx is small.
According to calculations made from Hiratsuka’s (1920) data the
exuviae of 5th instar larvae contain an average of 0.56 mg N, or about
0.54% of the N eaten and 0.91% of the N digested during the instar.
However, the average N content of the silk in one cocoon is about 41 mg,
or about 35% of the N eaten and 58% of the N digested during larval
life (calculated from data of Hiratsuka, 1920). Balogh and Gere (1953)
confused thei: data by lumping the exuviae and silk with the feces.
Table IX shows that the E.C.I. (N) ranges from 12% (on a plant
which supported only poor growth) to 62%. Unfortunately, without
more data there is no way to tell just how much of this variation is due
to differences in the apparent digestibility of N and how much to differ-
ences in the amount of digested N which is metabolized rather than
stored. Only Hiratsuka (1920) supplied sufficient data to permit calcu-
lation of the corresponding corrected C.A.D. (N) and E.C.D. (N). His
data show that Bombyx incorporates almost all of the digested N (92%)
in its body (Table IX). The generally high C.A.D.’s (protein N) would
lead one to expect the C.A.D. (N) to be generally high. The following
C.A.D.’s (protein N) are available in addition to those listed in Table
IX: Agluis, 63% ;Malacosoma, 72% ;Pieris, 83% (calculated from data
of Evans, 1939b) and Prodeniu, 91% (Crowell, 1941). However, it
should be noted that the amount of protein N “digested” and the
amount absorbed are not necessarily the same. A difference could result
if, as seems likely, some of the peptides and amino acids resulting from
the breakdown of protein pass out in the feces. If these breakdown pro-
ducts of protein are not all accounted for, the “digestibility” of protein
N will be greater than its absorbability, and the digestibility of non-
protein N will appear to be less than it actually is. With most leaf-
feeding insects the amount of non-protein N utilized is likely to be
small since protein N has been found to make up generally from 80%
to over 90% of the total N in leaves (Evans, 1939a, b; Fraenkel, 1953;
Hiratsuka, 1920).
Recently McGinnis and Kasting (1966) used the chromic oxide
method to measure both dry matter and nitrogen utilization by 5th
instar nymphs of MeIanoplus bivittatus.
Table X shows that the digestibility of sugars is very high, the C.A.D.’s
for total sugars ranging from 59% to 94%. The non-reducing sugars,
essentially sucrose in leaves, appear to be highly digestible, but it does
not follow that their breakdown products have been completely ab-
sorbed. It is quite probable that a part of the hydrolysates of sucrose,
TABLE X
The coefficients of apparent digestibility of carbohydrates and lipids
Insect, food and authority Reducing Non-red uc i ng Total Starch Total sugars Lipids
sugars sugars sugars and starch
Aglais urticae
Urtica dioica leaves 50" 13' 65' 9' 38" 55"
Evans (1939b)
Bombyx rnori
Morus alba leaves - - - - 4? 60"
Hiratsuka (1920)
Cliilo suppressalis
Autoclaved rice stems 63 96 87 54 61 -
Malacosorna neustria
Salix viniinalis leaves 64" 88" 76" 35" 68" 27"
Evans (1939b)
Phalera bucephala
Corylus aoellana leaves 71" 93 78" 0 46' 63"
Evans (1939a)
Pieris brassicae
Young cabbage leaves 58" 62" 59" 0 29" -
Evans (1939b)
Prodenia eridania
Phaseolus vulgaris leaves 56 99 85" 0 65" -
Crowell (1941)
Tenebrio molitor
Wheat bran 53" 95" 94" 59 54" 73"
Evans and Goodliffe (1939)
The experiments in which these data were obtained are summarized in Table IV. Calculated from author's data.
CONSUMPTION A N D U T I L I Z A T I O N O F FOOD 277
the reducing sugars fructose and glucose, are passed out with the feces.
The presence of these hydrolysates in the feces would make it appear
that the reducing sugars are less digestible than they actually are.
The non-reducing sugars would presumably be hydrolysed very
rapidly by digestive enzymes. The fact that some of the non-reducing
sugars pass through the gut unhydrolysed suggests that a corresponding
part of the food was simply not reached by the digestive enzymes.
The digestion of starch varies widely (Table X). Among the leaf-
feeding species investigated only Malacosoma digested a large percent-
age of the starch. Aglais digested very little starch, and the three
remaining leaf-feeders digested no starch at all. Another leaf-feeder,
Automeris, has also been found not to digest starch(Brown, 1930).Brown
(1937b) concluded from an analysis of its feces that Melanoplus bivittatus
had not digested starch. He had not, however, measured the amount
of starch intake.
Both Tenebrio and Chilo normally eat foods with a high content of
starch, and both digested over 50% of the starch they ingested (Table X).
Ullman (1932) found that amylase from several species of insects,
including Tenebrio and several leaf-feeding Lepidoptera, did not hydro-
lyse whole unbroken grains of starch. However, starch was hydrolysed
after the outer layer of amylopectin had been broken by cooking the
starch or grinding it with sand. It would be interesting to know if Chilo
was able to digest a large percentage of starch in Hirano and Ishii's (1962)
experiment because the grains had been ruptured by autoclaving.
Tsutsui and Sato (1954) had previously reported that Chilo excretes
starch unaltered. They found that the starch is, however, gradually
hydrolysed to simple carbohydrates in the feces. They suggested the
hypothesis that Chilo may be coprophagous and thus utilize these starch
hydrolysates. Hirano and Ishii (1962) felt that their findings are in
accord with this hypothesis.
Hiratsuka (1920) reported that Bombyx larvae do not digest the crude
fiber fraction of mulberry leaves. Phaleru larvae do not digest cellulose
(Evans, 1939a). Available data suggest that on the whole leaf-feeding
insects do not digest cellulose (Trager, 1953; Wigglesworth, 1965).
Tenebrio, however, digests about 36% of the hemicellulose which it
ingests (Evans and Goodliffe, 1939). Some wood-feeding insects do, of
course, digest cellulose either with or without the aid of intestinal
symbionts.
With the exception of Malacosoma the apparent digestibility of lipids
is fairly high (Table X). Hiratsuka (1920) found the average E.C.T. of
lipids for the five instars of Bombyx to be about 213%. Thus Bombyx
larvae store over twice as much fat as they digest, indicating the trans-
formation of carbohydrates into fats.
The nutritive ratio,
% digestible carbohydrate + (% digestible fat x 2-25)
% digestible crude protein
has been used as an expression of nutrient balance for domestic animals.
Digestible fat is multiplied by 2.25 since its heat value is that much
greater than the heat value of carbohydrate. The nutritive ratio for
Bombyx rose from the 1st to the 5th instar, 1.16, 1.25, 1.21, 1.26 and
1.50, respectively. The comparatively wide ratio obtained for the 5th
instar, according to Hiratsuka (1920), reflects the greatly increased rate
of fat deposition in that instar. Evans (1939b) gave nutritive ratios of
0.65 for Malacosoma, 0.32 for Aglais, 0.44 for Pieris and 1-18 for
Phalera. He pointed out that these ratios differ widely from ratios rang-
ing from 4-7 to 7-0 determined for growing cattle, sheep and pigs. The
nutritive ratio for Tenebrio molitor, 4.7, appears to be exceptionally
wide for an insect, and probably reflects the oxidation of large quantities
of carbohydrate as a source of metabolic water (Evans and Goodliffe,
1939).
V I I . UTILIZATION OF E N E R G Y
The coeficient of metabolizable energy (C.M.E.), usually expressed as
a percentage, is calculated as :
gross energy in food eaten - gross energy in feces
C.M.E. =
gross energy in food eaten
The gross energy (or heat of combustion) of a substance is usually
measured with a bomb calorimeter and is defined as the energy liberated
as heat when the substance is completely oxidized to COz and H20
(Kleiber, 1961; Tyler, 1964). Metabolizable energy has been defined for
mammals as the gross energy in the food eaten less the gross energy in
the feces, urine and the methane produced in digestion. This is the energy
available for the production of heat, body substance or work (Kleiber,
1961). The above formula does not take into account the possibility that
insects produce methane or other digestive gases. Metabolizable energy
for insects as here defined is thus comparable to metabolizable energy for
birds as defined by Kendeigh (1967, personal communication).Hiratsuka
(1920) subtracted the gross energy in the feces (urine included) from the
gross energy of the food eaten by Bornbyx and called this value the
physiologically available energy. He made no mention of the possibility
that insects may produce digestive gases. I have found neither qualitative
nor quantitative data on the production of digestive gases in insects.
Krogh (1916) stated that carnivorous animals usually produce small
quantities of methane, but that herbivores may produce large quantities.
The eficiency of storage of ingested energy is calculated as:
gross energy stored in body
E.S.I. (E) =
gross energy in food eaten
The eficiency of storage of metabolizable energy is calculated as :
gross energy stored in body
E.S.M. (E) = x 100
gross energy in food eaten - gross energy in feces
Hiratsuka’s (1920) study of Bombyx remains to this day the most
complete study of the intake and expenditure of energy by an insect.
TABLE XI
Comparison of the utilization of dry matter and energy by Bombyx niori
To end of 5th instar To newly emerged adult

Dry matter Energy Dry matter Energy
- _____
A.D. 37 C.M.E. 4 2 A.D. 37 C.M.E. 42
E.C.I. 23 E.S.I.(E) 28 E.C.I. 8 E.S.I.(E) 12
E.C.D. 62 E.S.M.(E) 67 E.C.D. 22 E.S.M.(E) 28
Conversion indices are based either on gross gain from hatching to the 5th instar just
before spinning or on net gain from hatching to the newly emerged adult. Calculated from
data of Hiratsuka (1920).
Table XI shows that the utilization of energy is higher than the utiliza-
tion of dry matter, but that the two are roughly comparable. Hiratsuka
(1920) also found that the consumed portion of the leaves had a some-
what greater heat value than the total of leaves supplied to the larvae.
Table XI1 summarizes the utilization of energy by the five larval
instars of Bombyx. The C.M.E. and the E.S.I.(E) vary with age much as
do the corresponding A.D. and E.C.I. of dry matter (Tables VII and
VIII), declining in the early instars and then tending to remain more or
less constant. The E.S.M.(E), however, increases with age, reaching its
peak in the 5th instar, while the corresponding E.C.D. of dry matter
tends to remain relatively constant from instar to instar (see Section
HEAT AND WORK
280
037 kcol
d 69% 047 k c d
8 6% 0'12 kcol
.L lnm70 FULLY FED LARVA PUPA ADULT

2.00 hcol. 370% k c d , 27.47. -DEATH
3 5 3 kcol. 652.1.
,
\
PUP01
h
Y Exwoe Spermotozoo
G. P. W A L D B A U E R
Feces a Urine Silk 005kcal.09%
7 7 2 hcd 1.13 k c o 1 . 2 0 9 %
ENERGY IN PRODUCTS
A HEAT AND WORK
Q 061 hcol
0l5kcol
ADULT -DEATH
3 hcal. 28.5%
Feces B urine
8.76 kcol First 4 Exuvme
0'03 h c d , 0 4 %
ENERGY IN PRODUCTS
Fro. 1. The energy economy of male and female Bombyx mori (based on data of Hiratsuka, 1920).
CONSUMPTION A N D UTILIZATION O F FOOD 28 1
XI1
TABLE
The utilization of energy by the larval instars of Bombyx mori
Instar C.M.E. E.S.1.Q E.S.M.(E)

I 52 32 61
I1 49 29 60
m 42 28 66
N 44 29 66
V 42 28 68
Conversion indices are based on gross gain in each instar. Calculated from data of
Hiratsuka (1920).
VEl). The large amount of energy stored during the 5th instar is, of
course, needed to support the non-feeding pupa and adult.
Figure 1 is a graphic representation of Hiratsuka’s (1920) elucidation
of the energy economy of Bombyx. The female ingests about 16% more
energy and retains about 21% more metabolizable energy than the male.
Very little of the metabolizable energy, about 2%, is lost in the exuviae,
although about 20% is expended in the silk. From hatching to the com-
pletion of reproductive activity about 52% of the metabolizable energy
is respired by either sex. A large share of the metabolizable energy of the
female, over 12%, goes into the eggs. This is not much less than the
extra 17% (1.12 kcal) of metabolizable energy available to the female in
comparison with the male. After reproductive activity is completed the
female goes to her death retaining less than 14% (0-89kcal) of the total
metabolizable energy while the male goes to his death retaining over
25% (1.37 kcal).
Erhan et al. (1966) looked for circadian rhythms in the utilization of
energy by Bombyx. They measured the ingestion, excretion and metabol-
ism of energy for 24 hr in the 4th and 5th instars, but found no evidence
of circadian rhythms.
Kitzawa (1959) quoted Nakamura’s unpublished data on the utiliza-
tion of dry matter and energy by the larvae of Pieris rupae. The methods
used to obtain the data were not described. The accuracy of the data is
doubtful since the A.D. and C.M.E. for the 2nd instar, 95% and 97%,
respectively, are unbelievably high.
Teissier’s (1931) figures are stated to represent the amount of energy
or dry matter consumed by the larvae of Tenebrio. However, his method
actually yields the metabolizable energy or the approximate digestibility
of dry matter. He provided groups of larvae with a known dry weight of
282 G . P. W A L D B A U E R
food, and at the end of the experiment determined the dry weight of the
mixture of left over food and feces. His figures for “consumption”
represent the difference between these weights. The mean E.S.M.(E)
was 26% for larvae fed wheat flour and 32% for larvae fed powdered
milk (see also Table IV). However, the larvae were kept at conditions of
temperature and moisture far below the optimum and grew very
slowly. The wheat flour and the powdered milk contained only 12.5%
and 2.5% moisture, respectively. The experiments were run at ambient
temperatures of about 17°C and 20°C, respectively.
ACKNOWLEDGEMENT
I wish to thank my colleague, Dr. Gottfried Fraenkel, for his critical
reading of the manuscript and numerous helpful suggestions. Errors and
omissions, however, remain solely my responsibility.
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The Consumption and Utilization of Food by Insects

11. CONSUMPTION, GROWTH A N D U T I L I Z A T I O N INDICES

F = fresh or dry weight of food eaten

B . DIGESTIBILITY A N D EFFICIENCY OF CONVERSION

1. Conversion of ingested food

of the undigested food. The gases produced in digestion, mainly C 0 2

3. Conversion of digested food

111. MEASURING A N D UTILIZATION

Protoparce which attained a maximum fresh weight of 7.14 g (after

% wt loss of aliquot x wt food introduced2 + wt uneaten food

Range of % dry Mean difference in % dry matter of:

* From Waldbauer (1964).

exudation was sometimes somewhat less than the rate of honeydew

M F = concentration of marker in food

The chromic oxide technique was subsequently used by McMillian et

obtained by the chromic oxide method. Larvae fed a diet containing

S1= total sugars in ethanolic extract of a sample of flour

% dry C.I. calculated from :

* From Waldbauer (1964). Unpublished data of Waldbauer. Not normally eaten;

So0 Hoo and Fraenkel(l966) presented a number of dry weight C.I.'s

Insect, food and authority O; of total consumption by instars

A . LIMITATIONS OF THE DATA

Temp. (C) Sex and Instar

Temp. ((3 Sex and Instar

(Lepidoptera: Lymantriidae) ca 75% (Fagaceae) Instars V and VI 21 - -

Temp. (0 Sex and Instar

(Lepidoptera: Pieridae) - Brassica oleracea Part of (24 hr?) last instar

Temp. (C) Sex and Instar

30" Blood of man Instar I (unfed) to

or a few other species of Moraceae. Protoparce in nature eats tobacco or

Several comparisons of the utilization of different foods by the same

effect of crowding was the greatest on the nutritionally best plants.

4th instar 5th and 6th instars

a From Sattler (1939).

Little is known concerning the effecton food intake and utilization of

1. Variation with age

All figures are OR a dry weight basis unless otherwise indicated.

Indices are based on gross weight gain unless otherwise indicated.

female and male remained almost constant. In other species efficiency

Beet leaves - 31" 97" w

To end of 5th instar To newly emerged adult

.L lnm70 FULLY FED LARVA PUPA ADULT

A HEAT AND WORK

Instar C.M.E. E.S.1.Q E.S.M.(E)

Praha 1959, 317-320, Czechoslovak Acad. Sci., Prague.

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