Leaves of Populus tremuloides and Salix babylonica held in the laboratory as for feeding insects were analyzed for total nitrogen, total phenolics, and total sugars at 0, 3, and 6 days. More often than not, the three components varied due to leaf age, time since excision, or temperature. Temporal changes altered nutritional balance and probable impact on insect performance, Results suggest that other woody plant species would not remain nutritionally stable for long in the laboratory

Miller, William E

English

ValpoScholar

The Great Lakes Entomologist 
Volume 20 
Number 1 - Spring 1987 Number 1 - Spring 1987 Article 4 
April 1987 
Change in Nutritional Quality of Detached Aspen and Willow 
Foliage Used as Insect Food in the Laboratory 
William E. Miller 
University of Minnesota 
Follow this and additional works at: https://scholar.valpo.edu/tgle 
 Part of the Entomology Commons 
Recommended Citation 
Miller, William E. 1987. "Change in Nutritional Quality of Detached Aspen and Willow Foliage Used as 
Insect Food in the Laboratory," The Great Lakes Entomologist, vol 20 (1) 
Available at: https://scholar.valpo.edu/tgle/vol20/iss1/4 
This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. 
It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. 
For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 
1987 THE GREAT LAKES ENTOMOLOGIST 41 
CHANGE IN NUTRITIONAL QUALITY 01<' DETACHED ASPEN A D 

WILLOW FOLIAGE USED AS INSECT FOOD IN THE 

LABORATORY 

William E. Miller] 
ABSTRACT 
Leaves of Populus tremuloides and Sali.r: babylonica held in the laboratory as for 
feeding insects were analyzed for total nitrogen, total phenolics, and total sugars at 0, 3, 
and 6 days. More often than not, the three components varied due to leaf age, timc since 
excision, or temperature. Temporal changes altered nutritional balance and probable 
impact on insect performance, Results suggest that other woody plant species would not 
remain nutritionally stable for long in the laboratory, 
Although artificial diets are available for fecding many kinds of insects in the 
laboratory, natural foods may be used for convenience or for more purposeful reasons. 
Natural foods are oftcn uscd as controls in the evaluation of artificial diets (Singh 1977). 
For herbivores, foliage-bearing branch lets with cut ends usually in water are provided for 
stated periods or until consumed. While the nutritional quality of fresh foliage for insects 
is 
becoming better known, the stability of detached foliage 
is uncertain. 
Reported here are chemical analyses of foliage from detached aspen and willow 
branchlets held in the laboratory as for insect feeding, The components analyzed are 
important indicators of nutritional quality: total nitrogen, total phenolics, and total sugars. 
The dietary significance of these components for insects might be succinctly characterized 
as positive for nitrogen (Mattson 1980) and sugars (House 1974), and negative for 
phenolics (Levin 1971). 
MATERIALS AND METHODS 
Mid-crown branch tips 20-30 cm long were cut at mid-day from 4-m-tall quaking aspen 
(Populus tremuloides Michx,) at one site. Similar sizcd pendant branch tips reachable 
from the ground were cut from a mature weeping willow (Salix babylonica L.), Cut cnds 
were placed immediately in distilled water, and the branchlets promptly transferred to the 
laboratory for assignment to treatments. Cut ends were kept in distilled water throughout 
treatments. 
Treatments consisted of 12-h fluorescent light photophase and four temperature regimes 
in controlled environment chambers: (1) con tant 16cC, (2) constant 20°C, (3) day-night 
cycling 22-IOoC, and (4) day-night cycling 28-16°C, Aspen branchlets were held at the 
first two temperatures, and willow branch lets at the last two, At intervals of 0, 3, and 6 
days, 8-12 leaves with petioles were harvested from branch lets sampled along their 
length. Sampled branchlets were not returned to the experiment. 
Harvested leaves of each treatment were pooled in a Petri dish, and immediately stored 
at -15°C, The contents of each Petri dish were later freeze-dried for 45 h and ground in 
IDepartment of Entomology, University of Minnesota, SI. Paul, MN 55108. 
1
Miller: Change in Nutritional Quality of Detached Aspen and Willow Foliag
Published by ValpoScholar, 1987
42 THE GREAT LAKES ENTOMOLOGIST Vol. 20, No. 1 
a Wiley mill to pass a 40-mesh screen. Total nitrogen was measured by a micro- Kjeldahl 
method, total phenolics by photocolorimetry using Folin-Denis reagent (Rosenblatt and 
Peluso 1941), and total sugars by high-pressure liquid chromatography. 
Analyses of both species were carried out with branchlets of different ages: partly 
expanded leaves collected 16 May, and fully expanded leaves collected 10 June. On 
neither date was there noticeable natural insect or other injury to the foliage f either 
species. Foliage from the same sources in other years was used to feed insects in studies 
published (Miller 1977, Muggli and Miller 1980) and unpublished. 
RESULTS 
Nitrogen was the most stable component (Fig. 1). In aspen expanding (May) foliage, 
it ranged from 5.0 to 6.0%, apparently increasing at both temperatures at first, then 
leveling off. It persisted near 2.0% at both temperatures in expanded (June) foliage. In 
willow expanding (May) foliage, it ranged from 4.3 to 4.9%, and in expanded (June) 
foliage, from 3.6 to 3.8%, varying little with temperature regime. 
Phenolic content was nearly stable over the treatment period (Fig. 1). In aspen 
expanding (May) foliage, phenolic content ranged from 6.1 to 7.2%, and i  expanded 
(June) foliage, from 13.8 to 15.6%, where it appeared to increase at both temperatures. 
In willow expanding (May) foliage, it varied from 3.9 to 5.0%, and in expanded (June) 
foliage, from 4.7 to 7.5%. It appeared to increase in willow leaves at both temperature 
regimes. 
Sugar content declined steadily from 15 to less than II % in aspen expanding (May) 
foliage at both temperatures, and from 16 to less than 14% in expanded (June) foliage 
(Fig. 1). In willow foliage of both ages, sugar content increased from 16% at the lower 
temperature then declined, and at the higher temperature regime, it changed in the 
opposite way. 
DISCUSSION 
Because this study was replicated but once, sampling variation in components is largely 
unknown. Results are therefore preliminary. 
The large differences in total nitrogen and total phenolics between aspen expanding 
(May) and expanded (June) foliage (Fig. 1) conform with well known seasonal changes 
in these components (Mattson 1980). The lesser corresponding differences between 
willow expanding (May) and expanded (June) foliage are partly due to willow flushing a 
week or more earlier than aspen, and there being less difference in willow leaf expansion 
between collection dates. 
No pronounced trends related to temperature emerged for nitrogen in either species. For 
total phenolics, there may have been more accumulation in expanding (May) leaves of 
both species at lower than at higher temperatures, and the opposite in expanded (June) 
foliage. Temperature effects for total sugars were mostly nondescript, except for the 
divergence-convergence sequence in willow leaves (Fig. 1). 
Insects benefit in direct proportion t  foliar nitrogen content (Mattson 1980). Other 
things being equal, an increase in total nitrogen from 5.0 to 5.5 or 6.0%, as occurred in 
aspen expanding (May) foliage (Fig. 1), should benefit insects measurably. However, 
concurrent increase in phenolics can reduce nitrogen availability to insects (Mattson 
1980). 
The temporal increases in phenolics (Fig. 1) are similar to those following mechanical 
damage to intact foliage (Rhoades 1983). Results here supplement evidence of a localized 
phenolic response. Few dosage-response studies of phenolics and insects are available. 
Those reported show that for one-compound tests, amounts from 0.2 to 0.4% in artificial 
diets prolonged development, reduced pupal weights, and increased mortality compared 
with amounts from 0.1 to 0.2% (Shaver and Lukefahr 1969). Because phenolics appear 
2
The Great Lakes Entomologist, Vol. 20, No. 1 [1987], Art. 4
https://scholar.valpo.edu/tgle/vol20/iss1/4
1987 THE GREAT LAKES ENTOMOLOGIST 43 
Percent -------------, 
15.0 ~ June • __~-~-~ •
•--- I 
':::~ TOTAL iHENtOLICS ~ May _ _-0 60r~~~-------~--- ------ ~I .~:--- : 
3:0 [ . [~~:-- ______ 0 
0.0 21.0 ----'--___-'----__--l ~/ ,,~ I ~ TOTALSUGARS~ 
18.0 I June I ~ • ~~('oe/o .~ 111,,;- ,/ // I15.0:<~----___·~ t '~, ,.
'- ---0-_ '::Co::::-"", _NI~~ _.May '-,_,_~ ---- ---0 " .',,/ ____ 0 
12.0 lI ~! l '<0:'- - ~ 
::: r "'j ,-I__-'-__----'__---' 
0.0~-:--------.-: 
5.0.~ May IMay ._ r=-=i~o 
4.0 ~ June _ _ - - - -i - - - - - r
•I 
3.0 TOTAL NITROGEN 
June 
2.0 .-~~~.-- ­
.. ------020 C 
1.0 
.---.16 C ~-~---~L--~dW~ ~ ASPEN 
0.0 '---------'------'---­
o 2 4 60 2 4 6 
Time (days) 
Fig. 1. Dry weight percentages of three chemical components of expanding (May) and expanded 
(June) aspen and willow foliage on days 0, 3, and 6 at different temperatures in the laboratory. 
to act in a dosage-dependent manner, buildups as in aspen and willow expanded (June) 
foliage may adversely affect insect performance. However, undetermined facors like 
specific composition of the phenolics, and capacity of some insects to overcome their 
adverse effects, may be important. 
Although sugars are important feeding stimulants, their temporal fluctuation in 
detached aspen and willow foliage (Fig. 1) lies in a nutritionally significant range. 
3
Miller: Change in Nutritional Quality of Detached Aspen and Willow Foliag
Published by ValpoScholar, 1987
44 THE GREAT LAKES ENTOMOLOGIST Vol. 20, No.1 
Different insects have different qualitative and quantitative sugar requirements and 
tolerances (House 1974); despite such complexities, insects investigated by Beck (1957) 
and Matsuoka (1965) benefited in direct proportion to sugar concentrations between 
threshold and inhibitory levels. For another species investigated by Harvey (1974), 
change in sucrose concentration from 1 t  4% in artificial diets produced female pupae 
65% heavier in less time. If insects of similar constitutions consumed aspen and willow 
foliage at some combinations of leaf age and temperature in this study, they would 
encounter marked temporal changes in extractable energy. 
CONCLUSION 
Detached aspen and willow foliage not immediately consumed does not retain its 
original balance of nutritionally important components, although some components may 
persist at original levels under some conditions. Conditions of change in investigated 
components include age or de ree of leaf expansion, time, and temperature. The changes 
in detached leaves may not differ greatly from those on intact leaves of the source plants. 
Overall nutritional quality cannot be estimated for detached aspen and willow foliage 
because not enough is known about interactive or compensatory effects of changing 
components for specific insects. However, to avoid unknown and unwanted variation in 
insect perfomlance, investigators should discard detached foliage not consumed within a 
few 
hours. 
The similarity in changes observed here between aspen and willow foliage suggests that 
detached foliage 
of other woody plant species likewise does not remain nutritionally stable 
for long under insect rearing conditions in the laboratory. 
ACKNOWLEDGMENT 
Major support for this study was provided by the USDA Forest Service North Central 
Forest Experiment Station, St. Paul, MN. Total nitrogen was determined by Department 
of Soil Science Analytical Services, University of Minnesota; sugars by Ingman 
Laboratories, Inc., Minneapolis, MN. I thank S. R. Palmer for phenolic analyses, and L. 
S. Bauer for laboratory and other assistance. 
LITERATURE CITED 
Beck, S. D. 1957. The European com borer. Pyrausta nubialis (HObn.), and its principal host plant. 
IV. Larval saccharotrophism and host plant resistance. Ann. Entomo!. Soc. Amer. 50:247-250. 
Harvey, G. T. 1974. Nutritional studies of eastern spruce budworm (Lepidoptera: Tortricidae). 1. 
Soluble sugars. Canadian EntomoL 106:353-365.
House, H. L. 1974. Nutrition. pp. 1--62 in M. Rockstein (ed.). The physiology of Insecta, ed. 2. 
Academic Press, New York. 648 pp. 
Levin, D. A. 1971. Plant phenolics: An ecological perspective. Amer. Nat. 105:157-181. 
Matsuoka, M. 1965. Carbohydrate requirements of the eri-silkworm, Philosalllia cynthia ricini. 
Bull. Sericult. Exp. Sta. 19:499-516. 
Mattson, W. J. 1980. Herbivory in relation to plant nitrogen content. Ann. Rev. Ecol. Syst. 
11:119-161. 
Miller, W. E. 1977. Weights of Polia grandis pupae reared at two constant temperatures 
(Lepidoptera: Noctuidae). Great Lakes Entomol. 10:47-49. 
MuggJi, J. M., and W. E. Miller. 1980. Instar head widths, individual biomass, and development 
rate of forest tent caterpillar, Malacosollla disstria (Lepidoptera: Lasiocampidae), at two densities 
in the laboratory. Great Lakes Entomo!' 13:207-209. 
4
The Great Lakes Entomologist, Vol. 20, No. 1 [1987], Art. 4
https://scholar.valpo.edu/tgle/vol20/iss1/4
1987 THE GREAT LAKES ENTOMOLOGIST 45 
Rhoades, D. F. 1983. Herbivore population dynamics and plant chemistry. pp. 155-220 in R:F. 
Denno and M. S. McClure (eds.). Variable plants and herbivores in natural and managed systems. 
Academic Press, New York. 717 pp. 
Rosenblatt, M., and J. V. Peluso. 1941. Determination of tannins by photocolorimeter. J. Assoc. 
Offic. Agric. Chern. 24:170-181. 
Shaver, T. N., and M. J. Lukefahr. 1969. Effect of flavonoid pigments and gossypol on growth and 
development of the bollworm, tobacco budworm, and pink bollworm. J. Econ. Entomo1. 
62:643-646. 
Singh, P. 1977. Artificial diets for insects, mites, and spiders. Plenum, New York. 594 pp. 
5
Miller: Change in Nutritional Quality of Detached Aspen and Willow Foliag
Published by ValpoScholar, 1987


Change in Nutritional Quality of Detached Aspen and Willow Foliage Used as Insect Food in the Laboratory

Valparaiso University

The Great Lakes Entomologist Volume 20 Number 1 - Spring 1987 Number 1 - Spring 1987 Article 4 April 1987 Change in Nutritional Quality of Detached Aspen and Willow Foliage Used as Insect Food in the Laboratory William E. Miller University of Minnesota Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Miller, William E. 1987. "Change in Nutritional Quality of Detached Aspen and Willow Foliage Used as Insect Food in the Laboratory," The Great Lakes Entomologist, vol 20 (1) Available at: https://scholar.valpo.edu/tgle/vol20/iss1/4 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 1987 THE GREAT LAKES ENTOMOLOGIST 41 CHANGE IN NUTRITIONAL QUALITY 01<' DETACHED ASPEN A D WILLOW FOLIAGE USED AS INSECT FOOD IN THE LABORATORY William E. Miller] ABSTRACT Leaves of Populus tremuloides and Sali.r: babylonica held in the laboratory as for feeding insects were analyzed for total nitrogen, total phenolics, and total sugars at 0, 3, and 6 days. More often than not, the three components varied due to leaf age, timc since excision, or temperature. Temporal changes altered nutritional balance and probable impact on insect performance, Results suggest that other woody plant species would not remain nutritionally stable for long in the laboratory, Although artificial diets are available for fecding many kinds of insects in the laboratory, natural foods may be used for convenience or for more purposeful reasons. Natural foods are oftcn uscd as controls in the evaluation of artificial diets (Singh 1977). For herbivores, foliage-bearing branch lets with cut ends usually in water are provided for stated periods or until consumed. While the nutritional quality of fresh foliage for insects is becoming better known, the stability of detached foliage is uncertain. Reported here are chemical analyses of foliage from detached aspen and willow branchlets held in the laboratory as for insect feeding, The components analyzed are important indicators of nutritional quality: total nitrogen, total phenolics, and total sugars. The dietary significance of these components for insects might be succinctly characterized as positive for nitrogen (Mattson 1980) and sugars (House 1974), and negative for phenolics (Levin 1971). MATERIALS AND METHODS Mid-crown branch tips 20-30 cm long were cut at mid-day from 4-m-tall quaking aspen (Populus tremuloides Michx,) at one site. Similar sizcd pendant branch tips reachable from the ground were cut from a mature weeping willow (Salix babylonica L.), Cut cnds were placed immediately in distilled water, and the branchlets promptly transferred to the laboratory for assignment to treatments. Cut ends were kept in distilled water throughout treatments. Treatments consisted of 12-h fluorescent light photophase and four temperature regimes in controlled environment chambers: (1) con tant 16cC, (2) constant 20°C, (3) day-night cycling 22-IOoC, and (4) day-night cycling 28-16°C, Aspen branchlets were held at the first two temperatures, and willow branch lets at the last two, At intervals of 0, 3, and 6 days, 8-12 leaves with petioles were harvested from branch lets sampled along their length. Sampled branchlets were not returned to the experiment. Harvested leaves of each treatment were pooled in a Petri dish, and immediately stored at -15°C, The contents of each Petri dish were later freeze-dried for 45 h and ground in IDepartment of Entomology, University of Minnesota, SI. Paul, MN 55108. 1Miller: Change in Nutritional Quality of Detached Aspen and Willow FoliagPublished by ValpoScholar, 198742 THE GREAT LAKES ENTOMOLOGIST Vol. 20, No. 1 a Wiley mill to pass a 40-mesh screen. Total nitrogen was measured by a micro- Kjeldahl method, total phenolics by photocolorimetry using Folin-Denis reagent (Rosenblatt and Peluso 1941), and total sugars by high-pressure liquid chromatography. Analyses of both species were carried out with branchlets of different ages: partly expanded leaves collected 16 May, and fully expanded leaves collected 10 June. On neither date was there noticeable natural insect or other injury to the foliage f either species. Foliage from the same sources in other years was used to feed insects in studies published (Miller 1977, Muggli and Miller 1980) and unpublished. RESULTS Nitrogen was the most stable component (Fig. 1). In aspen expanding (May) foliage, it ranged from 5.0 to 6.0%, apparently increasing at both temperatures at first, then leveling off. It persisted near 2.0% at both temperatures in expanded (June) foliage. In willow expanding (May) foliage, it ranged from 4.3 to 4.9%, and in expanded (June) foliage, from 3.6 to 3.8%, varying little with temperature regime. Phenolic content was nearly stable over the treatment period (Fig. 1). In aspen expanding (May) foliage, phenolic content ranged from 6.1 to 7.2%, and i  expanded (June) foliage, from 13.8 to 15.6%, where it appeared to increase at both temperatures. In willow expanding (May) foliage, it varied from 3.9 to 5.0%, and in expanded (June) foliage, from 4.7 to 7.5%. It appeared to increase in willow leaves at both temperature regimes. Sugar content declined steadily from 15 to less than II % in aspen expanding (May) foliage at both temperatures, and from 16 to less than 14% in expanded (June) foliage (Fig. 1). In willow foliage of both ages, sugar content increased from 16% at the lower temperature then declined, and at the higher temperature regime, it changed in the opposite way. DISCUSSION Because this study was replicated but once, sampling variation in components is largely unknown. Results are therefore preliminary. The large differences in total nitrogen and total phenolics between aspen expanding (May) and expanded (June) foliage (Fig. 1) conform with well known seasonal changes in these components (Mattson 1980). The lesser corresponding differences between willow expanding (May) and expanded (June) foliage are partly due to willow flushing a week or more earlier than aspen, and there being less difference in willow leaf expansion between collection dates. No pronounced trends related to temperature emerged for nitrogen in either species. For total phenolics, there may have been more accumulation in expanding (May) leaves of both species at lower than at higher temperatures, and the opposite in expanded (June) foliage. Temperature effects for total sugars were mostly nondescript, except for the divergence-convergence sequence in willow leaves (Fig. 1). Insects benefit in direct proportion t  foliar nitrogen content (Mattson 1980). Other things being equal, an increase in total nitrogen from 5.0 to 5.5 or 6.0%, as occurred in aspen expanding (May) foliage (Fig. 1), should benefit insects measurably. However, concurrent increase in phenolics can reduce nitrogen availability to insects (Mattson 1980). The temporal increases in phenolics (Fig. 1) are similar to those following mechanical damage to intact foliage (Rhoades 1983). Results here supplement evidence of a localized phenolic response. Few dosage-response studies of phenolics and insects are available. Those reported show that for one-compound tests, amounts from 0.2 to 0.4% in artificial diets prolonged development, reduced pupal weights, and increased mortality compared with amounts from 0.1 to 0.2% (Shaver and Lukefahr 1969). Because phenolics appear 2The Great Lakes Entomologist, Vol. 20, No. 1 [1987], Art. 4https://scholar.valpo.edu/tgle/vol20/iss1/41987 THE GREAT LAKES ENTOMOLOGIST 43 Percent -------------, 15.0 ~ June • __~-~-~ ••--- I ':::~ TOTAL iHENtOLICS ~ May _ _-0 60r~~~-------~--- ------ ~I .~:--- : 3:0 [ . [~~:-- ______ 0 0.0 21.0 ----'--___-'----__--l ~/ ,,~ I ~ TOTALSUGARS~ 18.0 I June I ~ • ~~('oe/o .~ 111,,;- ,/ // I15.0:<~----___·~ t '~, ,.'- ---0-_ '::Co::::-"", _NI~~ _.May '-,_,_~ ---- ---0 " .',,/ ____ 0 12.0 lI ~! l '<0:'- - ~ ::: r "'j ,-I__-'-__----'__---' 0.0~-:--------.-: 5.0.~ May IMay ._ r=-=i~o 4.0 ~ June _ _ - - - -i - - - - - r•I 3.0 TOTAL NITROGEN June 2.0 .-~~~.-- ­.. ------020 C 1.0 .---.16 C ~-~---~L--~dW~ ~ ASPEN 0.0 '---------'------'---­o 2 4 60 2 4 6 Time (days) Fig. 1. Dry weight percentages of three chemical components of expanding (May) and expanded (June) aspen and willow foliage on days 0, 3, and 6 at different temperatures in the laboratory. to act in a dosage-dependent manner, buildups as in aspen and willow expanded (June) foliage may adversely affect insect performance. However, undetermined facors like specific composition of the phenolics, and capacity of some insects to overcome their adverse effects, may be important. Although sugars are important feeding stimulants, their temporal fluctuation in detached aspen and willow foliage (Fig. 1) lies in a nutritionally significant range. 3Miller: Change in Nutritional Quality of Detached Aspen and Willow FoliagPublished by ValpoScholar, 198744 THE GREAT LAKES ENTOMOLOGIST Vol. 20, No.1 Different insects have different qualitative and quantitative sugar requirements and tolerances (House 1974); despite such complexities, insects investigated by Beck (1957) and Matsuoka (1965) benefited in direct proportion to sugar concentrations between threshold and inhibitory levels. For another species investigated by Harvey (1974), change in sucrose concentration from 1 t  4% in artificial diets produced female pupae 65% heavier in less time. If insects of similar constitutions consumed aspen and willow foliage at some combinations of leaf age and temperature in this study, they would encounter marked temporal changes in extractable energy. CONCLUSION Detached aspen and willow foliage not immediately consumed does not retain its original balance of nutritionally important components, although some components may persist at original levels under some conditions. Conditions of change in investigated components include age or de ree of leaf expansion, time, and temperature. The changes in detached leaves may not differ greatly from those on intact leaves of the source plants. Overall nutritional quality cannot be estimated for detached aspen and willow foliage because not enough is known about interactive or compensatory effects of changing components for specific insects. However, to avoid unknown and unwanted variation in insect perfomlance, investigators should discard detached foliage not consumed within a few hours. The similarity in changes observed here between aspen and willow foliage suggests that detached foliage of other woody plant species likewise does not remain nutritionally stable for long under insect rearing conditions in the laboratory. ACKNOWLEDGMENT Major support for this study was provided by the USDA Forest Service North Central Forest Experiment Station, St. Paul, MN. Total nitrogen was determined by Department of Soil Science Analytical Services, University of Minnesota; sugars by Ingman Laboratories, Inc., Minneapolis, MN. I thank S. R. Palmer for phenolic analyses, and L. S. Bauer for laboratory and other assistance. LITERATURE CITED Beck, S. D. 1957. The European com borer. Pyrausta nubialis (HObn.), and its principal host plant. IV. Larval saccharotrophism and host plant resistance. Ann. Entomo!. Soc. Amer. 50:247-250. Harvey, G. T. 1974. Nutritional studies of eastern spruce budworm (Lepidoptera: Tortricidae). 1. Soluble sugars. Canadian EntomoL 106:353-365.House, H. L. 1974. Nutrition. pp. 1--62 in M. Rockstein (ed.). The physiology of Insecta, ed. 2. Academic Press, New York. 648 pp. Levin, D. A. 1971. Plant phenolics: An ecological perspective. Amer. Nat. 105:157-181. Matsuoka, M. 1965. Carbohydrate requirements of the eri-silkworm, Philosalllia cynthia ricini. Bull. Sericult. Exp. Sta. 19:499-516. Mattson, W. J. 1980. Herbivory in relation to plant nitrogen content. Ann. Rev. Ecol. Syst. 11:119-161. Miller, W. E. 1977. Weights of Polia grandis pupae reared at two constant temperatures (Lepidoptera: Noctuidae). Great Lakes Entomol. 10:47-49. MuggJi, J. M., and W. E. Miller. 1980. Instar head widths, individual biomass, and development rate of forest tent caterpillar, Malacosollla disstria (Lepidoptera: Lasiocampidae), at two densities in the laboratory. Great Lakes Entomo!' 13:207-209. 4The Great Lakes Entomologist, Vol. 20, No. 1 [1987], Art. 4https://scholar.valpo.edu/tgle/vol20/iss1/41987 THE GREAT LAKES ENTOMOLOGIST 45 Rhoades, D. F. 1983. Herbivore population dynamics and plant chemistry. pp. 155-220 in R:F. Denno and M. S. McClure (eds.). Variable plants and herbivores in natural and managed systems. Academic Press, New York. 717 pp. Rosenblatt, M., and J. V. Peluso. 1941. Determination of tannins by photocolorimeter. J. Assoc. Offic. Agric. Chern. 24:170-181. Shaver, T. N., and M. J. Lukefahr. 1969. Effect of flavonoid pigments and gossypol on growth and development of the bollworm, tobacco budworm, and pink bollworm. J. Econ. Entomo1. 62:643-646. Singh, P. 1977. Artificial diets for insects, mites, and spiders. Plenum, New York. 594 pp. 5Miller: Change in Nutritional Quality of Detached Aspen and Willow FoliagPublished by ValpoScholar, 1987

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Change in Nutritional Quality of Detached Aspen and Willow Foliage Used as Insect Food in the Laboratory

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