Original article
The effects of sublethal exposure to diazinon, carbaryl and resmethrin on longevity and foraging
in Apis mellifera L.
K. E. MacKenzie M. L. Winston
Department
ofBiological Sciences,
Center for PestManagement,
Simon FraserUniversity,
Burna-by, BC,
Canada VSA 1 S6(received 11-1-1988, accepted 31-7-1988)
Summary — Topical,
sublethalapplications
of threeinsecticides, diazinon, carbaryl
andresmethrin,
were
given
to workerhoney
bees(Apis
melliferaL.)
of two age groups, 0 and 14days.
Fornewly emerged workers, carbaryl
was the most hazardous in sublethal amounts,adversely affecting
bothlongevity
andforaging
age. Resmethrin was intermediate ineffect,
and diazinon the least hazard-ous. This is different from
mortality
studies in which the effects ofcarbaryl
and diazinon weresimilar,
and resmethrin the most toxic.For
14-day-old workers,
there wereonly
twostatistically significant
differencesaffecting foraging
in the sublethal
studies,
and these were not conclusive.However,
inmortality
studiescarbaryl
wasthe least
toxic,
diazinon intermediate and resmethrin the most toxicpesticide.
Newly emerged
workerhoney
bees were more sensitive topesticide
exposure, in both acute and sublethaleffects,
than older workers. In order to evaluatepesticide
hazards to thehoney bee,
bothlaboratory
and field tests should be used. A fieldbioassay involving
sublethal exposure effects onlongevity
andforaging
may be useful in thisregard.
Apis
mellifera—longevity
-foraging
- sublethal dose — insecticideRésumé — Action de doses sublétales de
diazinon,
decarbaryl
et de resméthrine sur la lon-gétivité
et lebutinage
de l’abeille(Apis
mellificaL.).
On a montré que depetites quantités
depesticides pouvaient
causer des eHets néfastes autant aux abeillesprises
individuellementqu’à
lacolonie dans son ensemble. Cette étude évalue l’effet de
l’exposition
sublétale auxpesticides
sur lalongévité
et lebutinage
des ouvrières de colonies standard deplein champ.
On a fait des
applications topiques
sublétales de 3insecticides,
lediazinon,
lecarbaryl
et laresméthrine,
sur des ouvrières d’abeilles de deux groupesd’âge;
0 et 14jours.
On a testé les troispesticides
sur les abeilles naissantes et seulement le diazinon et lecarbaryl
sur les abeillesâgées
de
14 jours.
On a diluéchaque pesticide
dans l’acétone pour obtenir des taux de mortalité d’environ 5, 10 et 25%, telsqu’ils
sont déterminés dans les études de toxicitéaigüe.
Les ouvrières testées ontreçu 1 pl
de la solution choisie et les témoins 1ut
d’acétone. La survie des ouvrières a été contrôlée tous les 10 à 14jours.
On a observé tous les troisièmesjours
lebutinage
enbloquant
l’entrée de la ruche
pendant
30 minutes. Les abeilles testées trouvées hors de la ruche ont été classées comme butineuses.Les 3 colonies diffèrent
significativement
par lalongévité
etl’âge
des butineuses(Tableau 11).
Pour cette
raison,
dans lesanalyses
suivantes les 3 colonies ont été considéréesséparément.
Pour les ouvrières
naissantes,
lecarbaryl
est leplus
nocif aux dosessublétales,
affectant à la fois lalongévité
etl’âge
des butineuses(Tableau lit).
La resméthrine a un effet intermédiaire et le diazi-non est le moins nocif. Ces résultats sont différents de ceux obtenus avec la toxicité
aigüe,
pourlaquelle
lecarbaryl
et le diazinonproduiseni
des effets semblables et la resméthrine est laplus toxique (Tableau I).
Chez les ouvrières
âgées
de 14jours,
on n’a trouvé que 2 différencesstatistiquement significa-
tives pour les doses
sublétales,
mais elles ne sont pas concluantes(Tableau IV). Toutefois,
en cequi
concerne la toxicitéaigüe,
lacarbaryl
s’est montré le moinstoxique,
le diazinon d’effet intermé- diaire et la resméthrine laplus toxique (Tableau 1).
Les relations entre les 3 insecticides ne sont pas les mêmes dans les tests de toxicité
aigüe
etdans les tests des effets sublétaux. La
resméthrine, qui
a la toxicitéaigüe
laplus élevée, produit
des effets sublétaux moindres que ceux du
carbaryl, qui
a la toxicitéaigüe
laplus
faible. Les ouvrières naissantes sontplus
sensibles auxpesticides
que les ouvrièresplus âgées,
tant dans leseffets sublétaux
qu’aigus.
Afin d’évaluer la toxicité despesticides
pour lesabeilles,
il est nécessaire de faire des tests à la fois au laboratoire et sur le terrain. A cettefin,
un testbiologique prenant
encompte
les effets d’uneexposition
sublétale sur lalongévité
et lebutinage
serait utile.Apis
mellifica -longévité
-butinage
- dose sublétale - insecticideZusammenfassung — Wirkung
subletaler Dosen vonDiazinon, Carbaryl
und Resmethrin aufLanglebigkeit
und Sammelaktivität bei derHonigbiene (Apis
melliferaL.).
Auchgeringe
Men-gen von Pestiziden
zeigen
oft zerstörerischeWirkungen
sowohl auf die einzelne Biene als auch auf die Kolonie als Ganzes. In dieserUntersuchung
sollte der Einfluß von subletalen Dosen von Pesti- ziden auf dieLanglebigkeit
und das Sammelverhalten von Arbeitsbienen in Standard-Freilandvöl- kern beurteilt werden.Arbeitsbienen
(Apis mellifera)
aus zwei verschiedenenAltersgruppen (0
und 14Tage alt)
wurden mit subletalen Dosen von 3 Insektiziden
(Diazinon, Carbaryl
undResmethrin) topikal
behandelt. Alle drei Chemikalien wurden bei frisch
geschlüpften
Arbeitshienengetestet,
aber nurDiazinon und
Carbaryl
bei den 14Tage
alten Bienen. Jedes Insektizid wurden in Acetonaufgelöst,
und zwar so, daß Mortalitätsstufen von
ungefähr 5%,
10°/ und 25%vorgegeben
waren, die ausUntersuchungen
der akuten Toxizität bestimmt wurden. Die Testhienen erhielten1 pl der jeweiligen
Konzentration und die Kontrollbienen
1 pl Aceton.
DieÜberlebensrate
der Arbeiterinnen wurde 10-14täglich
kontrolliert.An jedem
drittenTag
wurde die Sammelaktivitätprotokolliert.
Dazu wurde dasFlugloch
für 30 mingeschlossen. Testhienen,
die sich außerhalb des Stockeseinfanden,
wurden als Sammlerinnengezählt.
Die drei in diesem Versuch verwendeten Kolonien unterschieden sich
signifikant
in derLangle- bigkeit
und im Sammelalter(Tab. 11).
Dehalb wurden die drei Völker bei denfolgenden Analysen separat
betrachtet. Bei frischgeschlüpften
Bienen warCarbaryl
in subletalen Dosen dasgefähr-
lichste Gitt. Es wirkte sich
nachteilig
sowohl auf dieLanglebigkeit
als auch auf das Sammelalter aus(Tab. III).
Resmethrin war im Effekt einmittelmäßiges
und Diazinon das amwenigsten gefährliche
Gitt. Dies war bei den Tests über die akute
Giftwirkung anders,
dort warenCarbaryl
und Diazinongleich
und Resmethrin dasgefährlichste
Gift(Tab. I).
Bei 14
Tage
alten Bienenergab
dieUntersuchung
derWirkung
von subletalen Dosen nur zwei statistischsignifikante
Unterschiede und diese waren nichtschlüssig (Tab. IV).
Jedoch war bei derUntersuchung
der akutenWirkung Carbaryl
das amwenigsten gittige,
Diazinon einmittelmäßiges
und Resmethrin das
gittigste Agens (Tab. 1).
Das Verhältnis der drei Insektizide bei der
Untersuchung
der akutenGiftwirkung
war also ver-schieden vom Verhältnis bei subletalen Dosen.
Resmethrin,
das am meisten akutgittige,
hatteweniger nachteilige
Effekte alsCarbaryl,
das amwenigsten
akutgittige.
Frischgesc h lüptte
Bienenwaren
empfindlicher
gegen Pestizide als ältereBienen,
sowohl bei akuten wie bei subletalen Gitt-wirkungen.
Um die Gefährlichkeit von Pestiziden zubeurteilen,
sollten daher Labor- und Feldver- suchedurchgeführt
werden. EinBioassay
zurUntersuchung
der Effekte subletaler Dosen auf dieLanglebigkeit
und das Sammelverhalten wäre sinnvoll.Apis
mellifera -Langlebigkeit
- Sammelaktivität - subletale Dosen — InsektizideIntroduction
The honey
bee(Apis mellifera L.)
isrecognized
as our mostimportant crop pollinator (McGregor, 1976; NRCC, 1981;
Crane
andWalker, 1983). North
Americanagricultural products valued
at some$
20billion
(U.S.) depend
in someway
on thehoney bee (Levin, 1984; Winston and Scott, 1984). Pesticide
usehas also be-
come an
essential part of modern agricul-
ture,
and thehoney
bee issusceptible
tomany commonly used chemicals (Ander-
son
and Atkins, 1968; Atkins, 1975;
Johansen, 1977, 1979, 1983; NRCC, 1981 It
isimportant, therefore,
to under-stand
how toprotect
thispollinator
frompesticide exposure and
toevaluate the hazards different chemicals
pose.A number of
different factors should
beconsidered when pesticides
areevaluated
for their hazard
tothe honey bee. Acute toxicity studies which
assessmortality
have
beenused almost exclusively
inthese considerations. However, exposure
tosublethal concentrations of pesticides
in the
field also could
have adverseeffects
oncolony
success,honey produc-
tion and crop pollination (NRCC, 1981).
Chronic feeding of such commonly used pesticides
asacephate (Stoner
etal., 1985), carbaryl (Winterlin and Walker, 1973),
dimethoate(Waller
andBarker, 1979; Waller
etal., 1979; Barker
etal., 1980; Stoner
etal., 1983) and carbofuran
(Stoner
etal., 1982)
towhole colonies
reducedhoney production,
broodrearing
and worker population size. Sublethal topical
treatments toindividual workers
resulted inimpairment
in the dance lan-guage with methyl-parathion (Schricker
and Stephen, 1970), reduced lifespan with
malathion or diazinon
(Smirle
etal., 1984;
MacKenzie, 1986); and altered foraging patterns with diazinon (MacKenzie, 1986)
and permethrin (Cox and Wilson, 1984). It
is important therefore
toevaluate both
acute
toxicity
and sublethal effectsof
apesticide
whengauging
its hazard tothe
honey
bee.A bioassay using longevity
as a meas-ure to
evaluate sublethal pesticide
expo-sure to the
honey bee has been sugges- ted (Smirle
etaL, 1984). This study
wasdesigned
to examine the sublethaleffects
ofthree insecticides,
and toevaluate the effect of
treatmentage
onthe results.
Inaddition
tolongevity, foraging
wasalso studied, since increasing foraging activity
and earlier foraging age have both been correlated with reduced longevity (Free
and
Spencer-Booth, 1959; Sekiguchi and Sakagami, 1966;
Winstonand Katz, 1981, 1982;
Winston andFergusson, 1985).
Inclusion of this task, foraging, may
increasethe sensitivity of the bioassay.
Materials and Methods
Chemicals
Commercial formulations of the three insecti- cides were used. The
organophosphorous
insecticide diazinon was formulated as a 12.5%
emulsifiable concentrate
(Later’s Diazinon,
PCP Act. No.11437,
Later ChemicalsLtd., Richmond, BC),
the carbamatecarbaryl
as a22.5% concentrate
(Wilson Liquid
Sevin Car-baryl insecticide,
PCP Act. No.17971,
Wilson LaboratoriesInc., Dundas, Ontario),
and thepyrethroid
resmethrin as a 0.25% solution(House
Plant InsectKiller,
PCP Act. No.16219,
Later ChemicalsLtd., Richmond, BC).
The for-mulations were diluted in acetone to the
requir-
ed concentrations. The formulation of
carbaryl
was first diluted
(1:10)
with distilled water tofacilitate further
mixing
with acetone.The colonies
Test colonies were located at Simon Fraser
University, Burnaby,
BC and studies conducted from June toSeptember
1985. Workers to betreated
were obtained from asingle colony (not
one of the
experimental colonies)
to minimizegenetic
variation. Combscontaining emerging
workers were
placed overnight
in an incubatorat 34°C and 50—70% RH.
Newly emerged
workers were marked on the dorsal surface of the thorax with coloured and numberedplastic
labels
(Opalithplattchen,
Chr.Craze, KG, Endersbach, FRG),
and either treated immed-iately
or introduced into a test hive and remov-ed and treated at the desired age.
Three colonies in standard
Langstroth deep equipment
were used. These colonies consis- ted of 2 boxes(supers)
with 10 frameseach, enough
workers to cover most of theframes,
and ahealthy laying
queen. A third super was added 10July
to alleviatecolony crowding.
This super was removed on 16
August
forhoney
extraction.Experimentation began
18June and continued until 5
September
for col-onies 1 and
3,
and 1August
forcolony
2.Queen
problems
occured in colonies 1 and 3.Colony
1 was found to bequeenless
on 10 0August
and wasrequeened
on 16August.
Avirgin
queen was found and removed fromcolony
3 on 27 June.Subsequently,
theoriginal
queen
appeared
to belaying normally,
but wassuperseded
sometime before 18July.
Treatments
Honey
bee workers were treated with atopical
insecticide
application
on the dorsal surface of their abdomens. An SMIMicro/Peftor-A O
wasused to
dispense
1pl
treatment solution toeach insect. Workers were held
by
the hindleg
with
forceps
while the insecticide wasapplied.
A group of workers were treated with 1
wl
acet-one as a control in all
experiments.
Concentra-tions
causing approximately
5%, 10% and 25%mortalities were used for each
pesticide
andeach age group in the sublethal
experiments.
Acute
toxicity
tests(LD 50 )
for the 3 chemi-cals were conducted to choose the dose range for sublethal exposure. Between 40 to 50 work- ers were treated at each of the 4 to 7 doses in the
mortality
tests.Newly emerged
workers(0 days)
were treated andplaced
intoholding
cages, while the older age group
(14 days)
wasmarked at emergence,
placed
into standardcolonies,
removed at theappropriate
age(grasped by
their hindlegs
withforceps
andplaced
intocages)
immobilized with carbon dioxide andtreated,
andplaced
intoholding
cages.
Sugar
syrup(50%)
and water were sup-plied
ad libitum viagravity feeding
bottles. Mor-tality
counts were taken at 24 h and data anal-yzed following
the WHO for insecticidesusceptibility
tests(Swarrop, 1966).
The result- antregression
lines were used to calculatedoses
corresponding
to the desiredmortality
levels.
For
experimental
purposes,newly emerged honey
bee workers(0 days
ofage)
were treat-ed with all 3
pesticides,
while14-day
old work-ers were treated with diazinon and
carbaryl only,
due to time constraints. This left a number of untreatedworkers, exposed
to neitherpest-
icide nor acetone, in each of the 3 colonies.Newly emerged
bees were treated on 18—20 June with the desireddosage
ofpesticide,
andplaced
in the hivethrough
thefeeding
hole inthe
top
board. Treatments for the14-day
oldbees were carried out on 3
July
for colonies 1 and3,
and 4July
forcolony
2. The hives weretaken
apart
and frames examined oneby
one.Marked workers were
picked
upby
the hindleg, treated,
held in a cage until all treatments for the hive werecompleted,
and reintroducedthrough
thefeeding
hole. The number of wor-kers treated per dose
ranged
from 50-65.Observations
Entrance observations
began
on 22 June andcontinued every third
day
until most of the beeswere dead. Entrances were blocked with wire
screen for 15
min, preventing
any workers fromleaving
orentering,
then the numbers of mark- ed workers at the entrance were recorded for another 15 min. Allreturning
workers wereconsidered to be
foragers (Winston
andKatz, 1982). Every
10—14days
the colonies werechecked for
surviving
workersby removing
andexamining carefully
each frame andrecording
all marked workers.
Data
analyses
Acute
toxicity
data wereanalyzed using
theWHO method for insecticide
susceptibility
tests(Swaroop, 1966).
The criterion ofnonoverlap- ping
95% confidence limits was used to deter- minesignificant
differences betweenLD 50 s
foreach age group.
For sublethal
studies, longevity
and the firstday
and duration offoraging
wereanalyzed.
Longevity
indays
was determined as the mid-point
between the lastday
seen and the subse-quent day survivorship
was examined.Forag- ing
was considered to commence on the firstday
a worker was seen outside the hive in the entrance observations. Duration offoraging
was the number of
days
between the first and last time a worker was seen outside the hive.The treated controls and the 3
dosages
of eachpesticide
in each age groupinitially
were com-pared by
one-wayanalysis
of variance and Duncan’smultiple
range test(Zar, 1984).
Thento examine the effects of different
pesticides,
allpesticide
treatments in one age group wereanalyzed by two-way analysis
of variance(i.e., pesticide
anddosage)
and the Student—New—man—Keuls
multiple comparison
test(Zar,
1984).
,To evaluate the
possible
effects of differ- ences incolony conditions,
data from the mark-ed,
untreated workers in the colonies werecompared by
one-wayanalysis
of variance and Student-Newman-Keulsmultiple comparison
test
(Zar, 1984)
for each of the 3categories
examined.
Results
Acute toxicity
tests of the 3chemicals
tonewly emerged workers
showedcarbaryl
and
diazinon
to be of similartoxicity
and
resmethrinthe
moreacutely toxic (Table I). The relationships between the
3chemicals for the 14-day old workers
were :
carbaryl
wasthe least toxic pest- icide, resmethrin the most, with diazinon
inbetween. In addition, all chemicals
were found tobe significantly
moreacutely
toxic to
newly emerged workers
than 14-day old
workers.Marked, untreated workers in the
3 test colonies weresignificantly
different in lon-gevity
and thefirst day of foraging (Table II). Colonies
1and
2 weresimilar
induration of foraging, but statistically differ-
ent from
colony
3.Therefore,
furtheranal-
yses
of sublethal
data werecarried
outseparately for the
3colonies.
Sublethal exposure
tocarbaryl
hadadverse effects
onlongevity and foraging
in newly emerged workers (Table 111). In
colonies
1and 2, control workers lived
longer than those treated with carbaryl,
and
began foraging
later incolony
1 andearlier
incolony
2.In colony 3, the only significant difference between
controland pesticide-treated bees
was thatbees treated with carbaryl LD 25 continued fora-
ging longer than either
of theother
2dosages
orthe control.
When the toxicities of
all 3pesticides
were
compared for newly emerged work-
ers,
carbaryl
caused moreadverse effects than the other
twochemicals, with
ageneral pattern of decreased longevity in
all
3colonies, and the latest foraging in colony
1(Table 111). Longevity
wasinter-
mediate in bees treated with resmethrin, and longer in those
treatedwith diazinon.
There were no
effects
onforaging
due toresmethrin
ordiazinon
treatment.When workers
weretreated
at 14days
of age with carbaryl
ordiazinon, there
were 2
statistically different results (Table IV). The control group continued foraging longer than any carbaryl
treatmentin col-
ony
2.In colony 3, foraging began earliest
in those workers
treated with diazinon
LD 5
, latest in those treated with carbaryl LD
5
, and
wasintermediate
in the other treatmentgroups.
Discussion
This
paperhas
evaluated bothlethal and sublethal effects of
3different insecticides
on
the honey bee. Carbaryl
wasfound
tobe
more harmful tonewly emerged work-
ers
than diazinon
orresmethrin in sub- lethal tests, although
resmethrin wasthe
mostacutely toxic pesticide. There
werefew
effects on14-day old workers in sublethal experiments; but
onceagain,
inacute tests
resmethrin
was the mosttoxic,
while
carbaryl
was the least toxicinsect- icide. These toxicity rankings
arethe
same as
those previously published (NRCC, 1981), and all
3chemicals have been placed into groups
withhigh
hazardto
the honey bee (Johansen, 1979; Crane and Walker, 1983).
In
rating pesticide hazard
tohoney bees,
acutetoxicity values of mortality
areoften relied
on(Anderson and Atkins, 1968; Atkins, 1975; Johansen, 1977;
NRCC, 1981),
evenwhen field exper- imentation is incorporated into
thehazard rating. The relationships between chem- icals
inlaboratory mortality and field stud- ies have been shown
tovary in
somecases.
For example, the synthetic pyre-
throids, such
aspermethrin, have high
acute toxicities in
mortality
testsand
areof
low hazard in the field due
tolow appli-
cation
rates andrepellent action (Pike et al., 1982; Shires et aL, 1984).
In
this study the relationships between
acute
toxicity and sublethal effects also
varied. Resmethrin,
the mostacutely toxic
compound,
wasonly intermediate
in itssublethal effects, while carbaryl, which
caused the most sublethal
effects
in thisstudy,
was notthe
mostacutely
toxic.Car- baryl has been found
tobe
aserious problem for beekeepers, causing high mortality
tofield colonies (Johansen
andBrown, 1972; Anderson and Glowa, 1984;
Melksham et al., 1985).
Instudies
on resi-dual toxicity
of 10pesticides, carbaryl
wasrated
asthe
mosthazardous
tohoney
bees due
toits persistence (Mansour
et tal., 1984; Mansour and Al-Jalili, 1985).
These
2characteristics, adverse effects of sublethal exposure
and aresidual nature, may
accountfor the beekeeping problems
caused by carbaryl. These results suggest that, in addition
tolaboratory and field experimentation, sublethal exposure
stud- iesshould be
usedwhen rating
thehazards of chemicals
tothe honey bee.
Foraging and longevity
areboth
cat-egories that have been shown
tobe affected by sublethal exposure
topesti-
cides (MacKenzie, 1986;
and thisstudy).
Foraging and longevity
arerelated, since
earlierforaging activity results
in reducedlifespan (Free and Spencer-Booth, 1959;
Sekiguchi and Sakagami, 1966; Winston
and Fergusson, 1985). Reduced longevity
was a more
consistent indicator
ofad-
verse
sublethal pesticide exposure than age of first foraging, but neither of these
categories
wereaffected
inall
cases.Considering
the number offactors, includ- ing worker activity, internal colony condi-
tions and forage availability (Maurizio, 1950; Woyke, 1984;
Winston andFergus-
son,
1985),
that caninfluence lifespan
and foraging in the honey bee, perhaps
this is
tobe expected. In
anearlier study,
Smirle
etal. (1984)
alsofound reduced
longevity
due tosublethal pesticide
exposure. Similar
results wereobtained due
tosingle sublethal exposure
todiazi-
non in
observation hives (MacKenzie, 1986). Probably other factors
are moreimportant
indetermining longevity and
regulating temporal division of labour than pesticide exposure, which
makes uncov-ering sublethal pesticide effects difficult.
Newly emerged
workers werefound
tobe
more sensitive topesticides, including
both acute
toxicity
andsublethal expo-
sure,than older workers. P,revious studies
on acute
toxicities agree with this result
(Koch, 1958/1959; Mayland and Burk- hardt, 1970; Smirle
etaL, 1984; Mac-
Kenzie, 1986). Differences in toxicities
couldbe related
to anumber
offactors, including differences
inenzyme levels
oractivities and absorptivity of the cuticle.
Other
insectshave been found
tobe
moresusceptible after moults and
atemerg-
ence
(Busvine, 1971). Enzymes
whichmetabolize xenobiotics are found
inthe
honey bee. Smirle
andWinston (1988)
found
that the concentrations
andacti-
vities of 2important
detoxificationenzymes, glutathione S-transferases and the polysubstrate monooxygenases,
werevery low
innewly emerged workers
com-pared
to anyother age from
1 to 7weeks.
This alone could
accountfor the great susceptibility of newly emerged workers
tovarious insecticides.
Stress
may
also have been a factor inthis study. Honey bees, especially newly emerged workers,
aresensitive
to stressin many forms such
asnarcosis (Beck-
mann,
1974; Ebadi et al., 1980),
sublethalpesticide exposure (Schricker and Ste- phen, 1970; Smirle
etal., 1984), and
increased activity (Lindauer, 1953; Free
andSpencer-Booth, 1959),
and neuralfunction has been shown
tobe adversely
affected by
some ofthese (Stephen and Schricker, 1970; Beckmann, 1974). As the
3
insecticides used
inthis study
areall
nervous
system poisons,
neuralimpair-
ment
may be involved
inboth the lethal and sublethal effects found.
This work suggests that
aninsecticide
has both
lethal andsublethal effects,
which do
notnecessarily
have the samerelationship for each chemical. In rating pesticide hazards
tothe honey bee,
abio- assay involving sublethal effects
shouldbe considered
inaddition
to acutetoxi- cities studies in the laboratory and the
field. Longevity and foraging appear
tobe
potential
measures to use inthe develop-
ment
of such
abioassay, although
morework remains to
be done
torefine this
technique. Evaluation of
awider range
ofpesticides, including herbicides
andfungi- cides, should
bemade,
and therelation- ship between
acute andsublethal pest- icide effects
onthe individual
andthe
colony studied.
Acknowlegments
We are
grateful
to S.Mitchell,
C.Scott,
J.Krul,
G.Dodgon,
S.Kolmes,
K.Ottaway,
E. Punnettand L.
Fergusson
forassistance;
to M. Smirle for advice onlaboratory procedures;
to the lateP. Oloffs and to S.
Kolmes,
B.Roitberg
and M.Smirle for review of the
manuscript,
and to G.Gries,
G.Tourigny
and A.Lesqueve
for thetranslations. This work has been
supported
inpart by
NRC Contract Research Grant No.5173 and an NSERC
operating grant
toM.L.W.,
and an NSERCpostgraduate
sholar-ship to K.E.M.
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