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Human Anatomy & Physiology (11th Edition)
11th Edition
ISBN:9780134580999
Author:Elaine N. Marieb, Katja N. Hoehn
Publisher:Elaine N. Marieb, Katja N. Hoehn
Chapter1: The Human Body: An Orientation
Section: Chapter Questions
Problem 1RQ: The correct sequence of levels forming the structural hierarchy is A. (a) organ, organ system,...
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Transcribed Image Text:Temperature-dependent
development and survival
of the lesser mealworm, Alphitobius diaperinus
L. M. RUEDA and R. C. AXTELL Department of Entomology, North Carolina State University.
Raleigh, North Carolina, U.S.A.
Abstract. Development, growth and survival of the lesser mealworm, Alphitobius
diaperinus (Panzer), were determined at six constant temperatures. No egg hatch or
larval development occurred at 17°C. At temperatures of 20, 25, 30, 35 and 38°C the
median development times (days), respectively, were for eggs (13.4, 6.0, 4.4, 2.6 and
2.6), larvae (133.0, 46.0, 26.2, 22.4 and 23.9), pupae (17.0, 8.0, 5.5, 4.0 and 4.1), and
from oviposition to adult emergence (164.4, 60.2, 37.9, 29.0 and 30.8). The Sharpe &
DeMichele (1977) model was used to describe the temperature-dependent development.
The mean egg survival (hatching) ranged from 61% to 86%, with lowest hatch at 20°C.
Survival of the larvae and pupae ranged from 32% to 73% and from 85% to 95%, respec-
tively, with lowest survival at 20°C. Pupae had significantly lower weights at 35°C and
adults at 38°C than at the other temperatures. Female pupae (20 mg) and female adults
(16 mg) were significantly heavier than male pupae (17 mg) and male adults (13 mg).
Adults (0.5-9 months old) laid 4-7 eggs per female per day at 25°C.
Key words. Alphitobius diaperinus, lesser mealworm, darkling beetle, temperature-
dependent development, poultry litter.
Introduction
The lesser mealworm, Alphitobius diaperinus (Panzer)
(Coleoptera: Tenebrionidae), is a major pest in commercial poul-
try houses (Axtell & Arends, 1990). Beetles are commonly found.
in the litter in turkey and broiler houses and in the accumulated
manure in caged-layer houses (Pfeiffer & Axtell, 1980; Safrit &
Axtell, 1984). The beetles harbour and potentially spread a wide
variety of viral, bacterial and fungal pathogens of poultry and
serve as intermediate hosts of cestodes parasitizing poultry
(Calnek et al., 1991; Despins et al., 1994; McAllister et al., 1994,
1995). Young birds feed readily on the beetle larvae and adults.
As a result of this feeding, there may be adverse effects on the
weight gains of the birds and enhanced pathogen transmission
(Despins & Axtell, 1994, 1995). The larvae often tunnel into
building insulation materials for pupation sites, and thus cause
structural damage and expensive repairs (Ichinose et al., 1980;
Safrit & Axtell, 1984; Vaughan et al., 1984; Turner, 1986). In
some cases, especially when infested litter or manure is spread
, on fields, the beetles move to nearby houses and businesses
causing a serious nuisance.
Information on the effects of temperature on the rates of de-
velopment and survival of various insect stages are necessary in
80
Correspondence: Dr R. C. Axtell, Department of Entomology, Box
7613, North Carolina State University, Raleigh, NC 27695-7613, U.S.A.
designing population and control strategy models. Data on the ef-
fects of temperature on the development of A.diaperinus (Preiss
& Davidson, 1968: Wilson & Miner, 1969; Ichinose et al., 1980)
have been reported. but are fragmentary and not provided in suffi-
cient detail for the mathematical calculations required in modern
temperature-dependent development models (Wagner et al., 1984).
This study was conducted to determine the developmental rates,
growth and survival of the immature stages of A.diaperinus un-
r several constant temperatures. Once the relationship between
temperature and immature development has been determined.
this information can be used to construct a simulation model to
predict the development of beetles under variable temperature
conditions. The temperature-dependent model of Sharpe &
DeMichele (1977) was used to describe developmental rates and
the distribution of development was described using the
approach of Stinner et al. (1975).
Materials and Methods
The eggs, larvae and pupae used in this study were F, progeny of
adult beetles collected in a turkey house in Duplin County, North
Carolina, U.S.A.. and maintained in moist turkey starter feed.
Larval rearing medium consisted of wheat bran (125 g), turkey
starter feed (40 g). brewer's yeast (3.7 g) and water (75 ml). The
wheat bran and feed were autoclaved prior to use. Laboratory
1996 Blackwell Science Ltd
![incubators with no light were used. Temperatures in the incuba-
tors were monitored continuously and the constant temperature
settings were maintained ±0.5°C. Removal of the insects from
the incubators at varying intervals for examinations at room
temperature was necessary but the time was kept to a minimum
(<10 min).
Egg development. The rates of development from egg deposi-
tion to hatch were determined using eggs deposited during a 4 h
period. The eggs were obtained by placing 100 beetles (1:1 sex
ratio; sex determined according to Barke & Davis, 1967) into
100 g of turkey feed mixed with 50 ml water in a screen-topped,
1 litre polyethylene container (12 cm diameter, 8.5 cm high). The
eggs were deposited on black construction paper in an assembly
consisting of ten pieces of paper (4 x 4 cm) held together at the
centre by a wire staple. Fifty eggs, still attached to the paper,
were placed in a polystyrene petri dish (6 cm diameter, 1.5 cm
deep) and five of these dishes were placed on top of a wire screen
in a 17 litre polyethylene tub (41 x 28 x 15 cm) containing 2 litres
of water. Relative humidity in each tub for each temperature was
measured using a hygrometer and ranged from 50% to 60%. A
tub was placed in each of the six incubators set at temperatures
of 17, 20, 25, 30, 35 and 38°C. At 12 h intervals, the eggs were
examined for hatching under a dissecting microscope, and
emerged larvae were counted and removed. In calculating egg
development rates it was assumed that all eggs were laid at the
mid-point of the 4 h adult oviposition interval and emerged at
the mid-point between the observations at 12 h intervals.
Larval and pupal development. Large numbers of eggs were
held at 3°C and those larvae hatching during a 6 h period were
used to determine development times at the various tempera-
tures. Larval development was assumed to begin at the mid-point
of the 6 h interval. The newly-hatched larvae were placed in 1
litre polyethylene containers (12 cm diameter, 8.5 cm high) con-
taining the rearing medium described above. Each container had
two 2.5 cm diameter holes covered with fine mesh in the side
and a cloth mesh top to allow air exchange. To prevent the me-
dium from drying, three rearing containers (one for each larval
density described below) were placed inside a 17 litre polyethyl-
ene tub (as described above) containing 2 litres of water.
The larvae were held at each of the six temperatures (17, 20,
25, 30, 35, 38°C) at larval densities of 1, 3 and 6 larvae per gram
of medium representing uncrowded, moderately crowded, and
crowded conditions, respectively. These densities were achieved
by placing 100 g, 33.3 g or 16.7 g of rearing medium in a rearing
unit and adding 100 newly hatched larvae. Rearing units were
measured at 24 h intervals to determine the stage of insect devel-
opment. As the larvae progressed from early instar to late instar,
the examination interval was reduced to as little as 6 h at the
higher temperature.
When a larva reached the prepupal stage (quiescent, slightly
curved body, non-feeding), it was removed from the larval rear-
ing unit and placed in a well (7 ml) ofa covered 12-well tissue
culture plate (No, 25815; Corning Company, Corning, New York,
U.S.A.). The well contained 1.00 g, 0.33 g or 0.17 g of rearing
medium corresponding to the three density levels at which the
larva had developed to the prepupa stage. The prepupae in the
tissue culture plates were returned to their respective tempera-
tures and subsequently were examined at 12 h intervals for the
presence of pupae and emergence of adult beetles.
Alphitobius diaperinus development 81
A sample of 50 pupae and 50 adults (25 males and 25 females
of each stage, $24 h old: sex determined according to Barke &
Davis, 1967) from each temperature and density combination
were weighed individually. Survival of larvae and pupae was
recorded.
Fecundity and longevity. The fecundity and longevity of adults
were determined at 25°C. 250 newly emerged adults (<24 h
old, 1:1 sex ratio) were placed into each of four 2 litre polyeth-
ylene containers with 200 g of rearing medium. When the beetles
were 2 weeks old, each container was initially examined to count
and remove dead adults. Thereafter, the medium was examined
monthly for 9 months and the dead adults counted and removed.
At the same time, the old medium was replaced with new me-
dium to remove any larvae and prevent the development of the
next generation of adult beetles. Errors in the numbers of dead
adults were probably small, even though it was possible that a
few dead adults were consumed by the live adults because no
partially eaten dead adults were found and no feeding on dead
beetles by live beetles was observed.
For fecundity measurements, adult beetles were transferred
from the containers to polystyrene petri dishes (5 females and 5
males per dish; five dishes per temperature). Each dish (6 cm
diameter, 1.5 cm high) contained 2 g of moist turkey brooder
feed and a paper assembly (as described above) for egg disposi-
tion. The eggs laid during an 8-day period were collected when
female adults were 0.5, 3, 6 and 9 months old. Adults for each
respective age were picked at random from the original beetle
stock. The paper assembly and the turkey feed were replaced
with new ones every other day during the 8-day ovipositon
period. Eggs laid on both paper assembly and turkey feed.
were counted; only about 2-3% of the eggs were in the feed.
Experimental design and Data analysis
A 3 x 5 factorial design (Sokal & Rohlf, 1981) was used for the
temperature-development experiments; factors included rearing
density at three levels and temperature at five levels (20.25.30.
35 and 38°C). There was no development at 17°C. Each beetle
rearing container was an experimental unit and each of the
fifteen combinations of temperature and larval density was
replicated four times. The differences in developmental rates of
larvae and pupae under various combinations of temperature
and density were tested using general linear model (GLM)
procedure (SAS Institute, 1988). Means were separated
using Tukey's test (P = 0.05).
The high temperature inhibition version of the Sharpe &
DeMichele (1977) development model was used to describe the
effect of constant temperature on median development rate (re-
ciprocal of median number of days to complete development).
This model with high temperature inhibition has the following
form (Equation 1):
r(K) =
RH025 x
K
298.15
xexp
HA 1
1.987 298.15 K (1)
1 + exp x
})]
where r(K) is the median development rate (days-¹) at
temperature K (Kelvin = °C+273). RH025, HA, TH and HH are
HH 1
1.987 TH](/v2/_next/image?url=https%3A%2F%2Fcontent.bartleby.com%2Fqna-images%2Fquestion%2F3b90a750-5b1d-4af1-8318-8922127502bb%2Fdd3623f2-d4d7-4808-a0b3-adf138882f26%2Fog2qza_processed.jpeg&w=3840&q=75)
Transcribed Image Text:incubators with no light were used. Temperatures in the incuba-
tors were monitored continuously and the constant temperature
settings were maintained ±0.5°C. Removal of the insects from
the incubators at varying intervals for examinations at room
temperature was necessary but the time was kept to a minimum
(<10 min).
Egg development. The rates of development from egg deposi-
tion to hatch were determined using eggs deposited during a 4 h
period. The eggs were obtained by placing 100 beetles (1:1 sex
ratio; sex determined according to Barke & Davis, 1967) into
100 g of turkey feed mixed with 50 ml water in a screen-topped,
1 litre polyethylene container (12 cm diameter, 8.5 cm high). The
eggs were deposited on black construction paper in an assembly
consisting of ten pieces of paper (4 x 4 cm) held together at the
centre by a wire staple. Fifty eggs, still attached to the paper,
were placed in a polystyrene petri dish (6 cm diameter, 1.5 cm
deep) and five of these dishes were placed on top of a wire screen
in a 17 litre polyethylene tub (41 x 28 x 15 cm) containing 2 litres
of water. Relative humidity in each tub for each temperature was
measured using a hygrometer and ranged from 50% to 60%. A
tub was placed in each of the six incubators set at temperatures
of 17, 20, 25, 30, 35 and 38°C. At 12 h intervals, the eggs were
examined for hatching under a dissecting microscope, and
emerged larvae were counted and removed. In calculating egg
development rates it was assumed that all eggs were laid at the
mid-point of the 4 h adult oviposition interval and emerged at
the mid-point between the observations at 12 h intervals.
Larval and pupal development. Large numbers of eggs were
held at 3°C and those larvae hatching during a 6 h period were
used to determine development times at the various tempera-
tures. Larval development was assumed to begin at the mid-point
of the 6 h interval. The newly-hatched larvae were placed in 1
litre polyethylene containers (12 cm diameter, 8.5 cm high) con-
taining the rearing medium described above. Each container had
two 2.5 cm diameter holes covered with fine mesh in the side
and a cloth mesh top to allow air exchange. To prevent the me-
dium from drying, three rearing containers (one for each larval
density described below) were placed inside a 17 litre polyethyl-
ene tub (as described above) containing 2 litres of water.
The larvae were held at each of the six temperatures (17, 20,
25, 30, 35, 38°C) at larval densities of 1, 3 and 6 larvae per gram
of medium representing uncrowded, moderately crowded, and
crowded conditions, respectively. These densities were achieved
by placing 100 g, 33.3 g or 16.7 g of rearing medium in a rearing
unit and adding 100 newly hatched larvae. Rearing units were
measured at 24 h intervals to determine the stage of insect devel-
opment. As the larvae progressed from early instar to late instar,
the examination interval was reduced to as little as 6 h at the
higher temperature.
When a larva reached the prepupal stage (quiescent, slightly
curved body, non-feeding), it was removed from the larval rear-
ing unit and placed in a well (7 ml) ofa covered 12-well tissue
culture plate (No, 25815; Corning Company, Corning, New York,
U.S.A.). The well contained 1.00 g, 0.33 g or 0.17 g of rearing
medium corresponding to the three density levels at which the
larva had developed to the prepupa stage. The prepupae in the
tissue culture plates were returned to their respective tempera-
tures and subsequently were examined at 12 h intervals for the
presence of pupae and emergence of adult beetles.
Alphitobius diaperinus development 81
A sample of 50 pupae and 50 adults (25 males and 25 females
of each stage, $24 h old: sex determined according to Barke &
Davis, 1967) from each temperature and density combination
were weighed individually. Survival of larvae and pupae was
recorded.
Fecundity and longevity. The fecundity and longevity of adults
were determined at 25°C. 250 newly emerged adults (<24 h
old, 1:1 sex ratio) were placed into each of four 2 litre polyeth-
ylene containers with 200 g of rearing medium. When the beetles
were 2 weeks old, each container was initially examined to count
and remove dead adults. Thereafter, the medium was examined
monthly for 9 months and the dead adults counted and removed.
At the same time, the old medium was replaced with new me-
dium to remove any larvae and prevent the development of the
next generation of adult beetles. Errors in the numbers of dead
adults were probably small, even though it was possible that a
few dead adults were consumed by the live adults because no
partially eaten dead adults were found and no feeding on dead
beetles by live beetles was observed.
For fecundity measurements, adult beetles were transferred
from the containers to polystyrene petri dishes (5 females and 5
males per dish; five dishes per temperature). Each dish (6 cm
diameter, 1.5 cm high) contained 2 g of moist turkey brooder
feed and a paper assembly (as described above) for egg disposi-
tion. The eggs laid during an 8-day period were collected when
female adults were 0.5, 3, 6 and 9 months old. Adults for each
respective age were picked at random from the original beetle
stock. The paper assembly and the turkey feed were replaced
with new ones every other day during the 8-day ovipositon
period. Eggs laid on both paper assembly and turkey feed.
were counted; only about 2-3% of the eggs were in the feed.
Experimental design and Data analysis
A 3 x 5 factorial design (Sokal & Rohlf, 1981) was used for the
temperature-development experiments; factors included rearing
density at three levels and temperature at five levels (20.25.30.
35 and 38°C). There was no development at 17°C. Each beetle
rearing container was an experimental unit and each of the
fifteen combinations of temperature and larval density was
replicated four times. The differences in developmental rates of
larvae and pupae under various combinations of temperature
and density were tested using general linear model (GLM)
procedure (SAS Institute, 1988). Means were separated
using Tukey's test (P = 0.05).
The high temperature inhibition version of the Sharpe &
DeMichele (1977) development model was used to describe the
effect of constant temperature on median development rate (re-
ciprocal of median number of days to complete development).
This model with high temperature inhibition has the following
form (Equation 1):
r(K) =
RH025 x
K
298.15
xexp
HA 1
1.987 298.15 K (1)
1 + exp x
})]
where r(K) is the median development rate (days-¹) at
temperature K (Kelvin = °C+273). RH025, HA, TH and HH are
HH 1
1.987 TH
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