The JapanSociety The Japan Society
of Medical Entomology andZoology and Zoology
ofMedicalEntomology
SANITARY
ZOOLOGY
as
al
s
tz 16
Vol.
No.4
m Studieson the behaviour'patternof feedingof three species americana cockroaches, Blattella germanica (L.), L.,and P. fuliginosa S.,with special reference to their responses to some constituents of rice bran and some carbohydrates
of
,Fleriplaneta
By
Hideakira Tsujii'
I. INTRODUCTION
those with
The American cockroach, PleTit)inneta amen'cana, the Smoky brown cockroach, P: faliginesa, and the Gerrnan eockroaeh, Blattella house-infesting roaches, germanicu, likeother are typically omnivorous They are similarly and meat fond of starchy materials preducts. Their omnivorous food preferencesare of interest in two ways. First,how theirbehaviour differs from that of oligoof feeding pattern one can phagous insects,and second, whether are strictly expect that the three species similar to each other in the patternor not. This paper deaTswith the results of some preliminary stu-
dieson
the responses
olfactory
and
of
these cockroaches
gustatorystimuli
II, MATERIALS
AND
to
species a
adults
were
constant
experiments
sects
did not they
prior
to
enough
of
old
the water
had
were
conditions
of sbown
that the test
leftunfed
with
three
solyents,
in-
suMciently
for several
They were duringthe experiments
test
Since the three species are rice bran, it was extracted
equipment
with
250C. Since prelimi-
to foodunless
respond
the three
of
nymphs
room
under
temperature
nary
starved,
or
made
supplied
days with
fond Soxhlet
especially
in
a
n-hexane,
ether
The n-hexane extract was furand basic ther fractionated intoacidic, neutra1, it was stearn-distillated as well fractions, and 1). The extract fractions S2, (Si, S3,A, (Fig. and
methanol.
fbodorigin
of
ne
METHODS
The insectswere bred at 27"C, 75% R H, 16 hrs. illuminationper day throughout
and
the year. The
were Food MF and NMF The behaviourexperiments "
nymphs ried
the American
of out
under
above-mentioned
Rat
"Orienta1
pe11ets
animal
used
with
as
newly
cockroach
their food
hatched were
conditions,
car-
but
Agricultural ChernicalsResearch Laboratories, Sankyo Co., Ltrl., Hiromachi,Shina-
1) S± ecop :
gawa-ku, Tokyo
Fig. 1 Fractlonating
schedule
of
rice
bran
extraets.
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of
(L'5,i)
B
and
the
N), the
(D) were
steam-distillate
biological assay. the
ments,
unds
Tn
effect
submitted
series
another
sugars
of
Rg), and forthe
and
and
of
related
experi-
feedingresponse of roaches was A more detaileddescription of each will be given, when necessary, in pages
methods
the results.
IIL ExpERIpt{ENTALREsuLTS
Rice bran Rs)
on
meter
A:
(Ro)and
pasted
were
up
and
old
the
icyinphs
residues
circles
to
inyice
(Ri,R2 2cm
of
and
in dia-
drawing paper (13cmX
Blattella germanica
Fig. 3 R3 leftunfed consumed
pot
the paper
and
starved
coclcroaches
box (12
cm
in
shelves)
was
are
that
rice
eat
were
responses
very
to
3). (Fig.
shelter
The
old
several
three
spe-
showed
to the three
),Tostef the bran direetlywithin a
rice
Ri
the results
attractive
After the
untill
wooden
hidden. box.
usually
of
not.
were
the adults of the residues Ri
leftuneaten
glass
complete
P, and
and
nymphs
americana
feedingof began
R2, but Rs was R2 were consumed of Blattella ger-
Fig. 2 Attractiyeness of rice bran for the three species, americana CNymphs) (Nymphs) B: Ptrriplaneta (Adults)C: PL .fitliginosa
untill
(R
bran
they
residues
came
rice to
the
bran was but the
minutes.
×
a
in depth)where
in Fig, 2. The
shown
few
12 cm
placed on the
Examples of
in
kept with a 9.5 cm, with
were ×
which
The paper cies
placed
was
in diameter, 25cm
tt18cm
spec:,es
rtysidues
in
a square-cut
13cm),
cockroaches
Responses of adults bi'anand itsextracted
1.
No. 4
X'c)].16
shelter
compo-
the
on
tested.
of
(Ri,R,
residues
・1) #tv l965"
'll
',?'i
Ri
amen'cana),
and
R2
were
Fig. 4 R2 fed when
used
alone
(B,germanica).
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of
!li Sb gtl 1965 :,lj' manica sively
the was
ate
R2
this
when
residue
ifTl
Vol.16
(257)
No, 4
exclu-
was
(Fig.4). PL .fitliginosaalso ate residues Ri, R2 and even R3 when rice bran for them (Fig. 5), This exnot available userl
carried out under a condition periment was identicalto that of Fig. 2 except that the paper was replaced by four small dishescontaining 100mg of rice bran and the three kinds of
Following the
respectively.
residues
the
ments,
the
of
weight
lastinstarnymphs
the
was
experi-
consumed
materials
by
determined.
I
Fig. 6
noo・.ij,'gfigg
feedingstimulative Si forthe
Attractive and
effects
of
adults
the of
extract
1'. americana.
a holein 1,ess than 30 minutes. activity of Si has Although the attractive been expected from the extinction of naturally the activity in the residue Ri, the results fur-
planetamade u:-・ge.,b
ther showed
that the three
speeies
both two fractions,one is the soluble in n-hexan and insolublein eat
is
other
in hours
Time
Fig. 5 Amount of rice bran cRol)and the ,IRt, residues R2 and Rs) consumed by P. .f)ttiginosa, All the
・the three
results
species
described above
indicatethat attracted to rice residues. How-
strongly
were
bran but scarcely ever,
rnore
to the three they ate the residues, preferring much Ri and R2 than R3 when they happened
to encounter 2,
these
residues.
Responses ofadults S2 and S3 The
extracts
・ether,and S3 with
Si
and
water
S2
dilutedwith
were
into 20
Si,
%
ooncentration
The dilutionswere dropped with pipets on a filter 15cm in diameter. paper Afterbeingdried,the impregnated filterpaper was placedin a glass pot which was identical in the experiment on rice bran to that used and the residues, The attitude of P. americana is shown in Fig. 6, The other two species The showed responses similar te P. americana ・extractSi was very attractive forthe:threespecles but S2 and S3 were not. The cockroaches ・carne to the Si spot straightly and devoured it ,together with the paper, the two species of Pleri-
in
volume
rate.
the
extract water,
residues
Si the
Ri,
R2, and slightly in Rs and is insolublein nhas two kinds hexane. Thus the formerfraction of characters, attractant and feedingstimulant, whereas the latterhas only one, feedingstimuIant. The main factorsof the latterfraction seem to be in the extracts S3 soluble in methaand ether, nol and insolublein both of n-hexane because the cockroaches fed on much more Ri they encountered these and R2 than R3 when residues.
3. to ex;traatfractions
largly in
contained
to
responded
Responses qf IV and B
adztlts
to extractf>uctions
A,
A, N The dilutions of the extract fractions way as described and B -'ere tested in the same on the extracts Si, S2 and Ss, though the ameunt of the fraction B was lessthan a tracial exlstence,
fractions were tested togeof B, gervnaniczxattracted at once were on the fraction N. After 24 hours, however, the both spots of N and A were found to have been bitten. When fractions A was tested alone, no cockroaches the fraction were within 30 minutes. The attracted at least When
ther, most
results
the three
the adults
of
indicate that in B. gexntanica activity of the extract Si islargely
seemingly
the attractive due to the neutral
fraction and
the
fdedingstimu-
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(258)
eskfu
lantactivity isattributable to both of the neutral and acidic fractions. The result was somewhat different in P. The adults amen'cana were americana. of R attracted at once to both of the spots N and A, and bit the spot A more heavilythan the spot N, so the attractive activity of the extract activity seems Si as well as the feeding stimulant to be due to both the fractions of N and A in theadults of this species. P, .faliginosashowed responses
to those
similar
newly hatchednymphs however, were attracted fractionN (Figs. 8, 9 and
The
4. Responses qf When
three species excited
of
and
D
were
not
/LE-ut.A2eEx:L+-oLs-E=Z
amentana, neutral
D
steam-distillate
ie
in the same fractions, the
tested extract
were
very
much
but not arThey scarcely bit the
to the paper
spots.
be
arrested
one
the
of
the
on
24 hours the spots were been bitten only a little. arrestant
reasons
o
why
5・
0
'
IP
3D
2P
Time in minutes
though
spots,
found to have
after
The poor
-moJe
amen'cana.
to the
cockroaches
this might
spots and
they
the
attracted
the
on
rested
4e
10),
dilutionwas
the
described on
as
"'ay
of only
to
adttlth'
P. P.
of
Vol. 16 No, 4
ig 19654
Attractiveness of the extraet Si activity of S3 for the newly hatcheclnymphs of R ainericana.
Fig. 7
and
arrestant
the steam-distil-
effect of
latemade
itdiMcultto determine its attractant kept, and the dishwas covered with a glass, in such a small effect glass pot as used here, The number of nymphs plate at once. gatherbecausethe cockroaches scarcely stayed on the The ing at each spot was periodically counted. The dithculty will be resolved by the spots. indicatesthat result isshown in Fig. 7, which addition of arrestant factorsto the distillate. the extract Si was attractive to the nymphs but This was proved in newly hatchednymphs of in the S3 not S3 was always not attractive when the extract S3 was usecl R americana other experiments even when used preliminary as an additional arrestant factor(Fig.11). alone. S3, however,was so effective in arresting the attracted nymphs that the double spot 5. Responses of' nE7e,(yr hatchedirymPhs qf'P. of
ainencana
Newly hatchednymphs
of
P. aineiicana
are
for the bioassaywhich needs a numand the attractant effect of individuals, materials is easily determinedeven in a of nymphs attracted dish by the number
suitable
ber the small
of
to the
spots.
ExtiactsSi
and
americana.
filterpaper tant spots
scarcely extracts
Si
and
S3 were
hatchedyoung nymphs of R were The dilutions dropped on a
(11cm in diameter),and of
Si, Ss
and
the
Si+Ss.
the greatestnum-
of
Si. The
one
at
the
extract
Si
spot
and
single
leastduring this
"'as
in
experiment
be eoncluded from the results that effect to・ Si has a strong attractant ot P. americana as・ newly hatchednymphs as to the adults, but its feedingstimulant
It may
the extract
resul-
the
After
well
being dried,the paper placed on papermade shelters in a petri-dish(15cm in diameter, 3em in depth) where 12・O nymphs were was
with
bittenboth in
the double
newly
were
crowded
was
the larger circle
S3
The dilutionsof the tested with
Si+S3
ber of nymphs. The arrestant effect of theexfound to be due・ tract S3 inthe double spot was to itsgreater effect inducing the nymphs to bite・ it,since the spot was bittenheavilyand the the smaller bittenspace was restricted within was 1ocatedin the center of' circle of S3 which
activity
was
as it was however,seems
phs
not
so enough
to the te be
adults,
to the young
The
highlyeffective
nym-
extract
S3.
in stimu-
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os ts th im latingthe feeding nymphs
and
Fractt'ons A The
acidic
action
of
both the
young
N
were
1965 lli VoL 16
adults
N A fraction
and
and
neutral
In the first experiment,
tested in similar the number of nymphs
ways.
observed
on
two
double
that on two the latters were single spots of S3, in the second by two doublespots of N+S3, and in replaced comthe third two doublespots of N+S3 were pared to two single spots of S3. The results in Figs. 8, 9 and 10. It is clear are shown that the fraction N was attractive forthe nymwhereas A was not, although the fraction phs A was attractive and feedingstimulative forthe
spots
A+S3
of
adults
of
(259)
No. 4
was
this species
compared
to
eacov=--'qem=nfix:igo-Bfi:z
(3).
1 eoamo=-:ove-nExq・g-o-0p:sz
Fig. 10 newly
Time in rninutes N for the Attractiveness of the fraction hatchednymphs of R ame7'icana. {2'1.
20
5
o
10
Time in minutes Fig. 8 Unattractiveness uf the fraction A for the newly hatchednymphs of
R
15e:n=-o
a.titericana.
le
eonno=-:
ge. ,,aS-o--21E,sz
c di=nfira:.-
5
c "-2SEOZ
o
o
5
'
IO
Time in minutes Fig. 9N Attractiveness of the fraction forthe newly hatchednymphs of R atnericana Cl).
o
o
.s
xe
Time in minutes Attractiveness of the
Fig. 11 for the newly
hatehednymphs
steam-dlstillate of
R
D
atnericana.
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di ts fu ig 1965 IF V'o]. 16 No. 4
Steam-distillate D The steam-distillate way
and
related
D
inthe
tested
was
same
found to be highlyattractive (Fig.
was
11). But the bittenspace was strictly restricted the S3 part in the double spots of D+ Ss. After being washed with 1% HCI and 3 was % NaOH, the distillate still attractive,
the
Responses qf
As the
already
methanol
group
the
of
chromatograms some
contain
and
several
ties.
adt{lts
to some
sttgars
and
cofnj)ounds
iJelated
described in the previous sections, soluble fractionS3 consists one feedingstirnulants. By thin layer the fraction was estimated to carbohydrates in largequantities
other
in
compounds
Therefbrethe
effect
of
Tab]e 1 Effectof different sugars compounds on the feedingof the a
R
g'.
and
and
related
ww
D-Arabinose
't
Xylese Ribose 1
---
Glucose
'
+
residues Ri R2 R3. In the firstexperiment 200mg
+
ff -+
ber of
from
paper and
placed in the
was
was
compound
r
"
111
Sorbitol Dulcitol
+
meso-Inositol
--
were
easily
The
results
Table 2 Effect of different melar
Mme
-
Sucrose Lactose
1
Raflinose
T!
-
Starch
Dextraii
1
One p]us sign by theatcockroaches, of
The sp"ts
number responded.
ancl
of
cl+'/
une
2
lm
z)V{・
r+-"
kriplaneta
L-Arabinose Galactose
-
2 M.
"
"
+
---
-
11-
-lr
1-
MaltoseSucrose
1411i'
110
-
j/i
l
-
-
+
+
+
"
1・1 '10 /r/-1-
Galactose Fructose pt(annitol Sorbitol pt{altoseSucrose
responded
the total
num-
1
T4
M.2
i
1・ I+ 1-
Ht-
/
respendecl
not (.-1i are
11,
' -"-+atnertcana -
1
/spot
signs
i4
i
loo
!
minus
in
substances
concentrations.
Tl-
L-Arabinese
i
+ .1e'xpresses
in Table 1
shown
1"
MaltoseSucrose
-
+
il
are
Jlerr )lanettz .fttliginosa
i
Pectin
nU.ber
ma -
rm
/
five times. by tufts of
tested
iO
L-Arabinose Galactose Fi'uctose Mannitol Sorbltol
/
Salicin Glycogen
Every
recognized
seven
100
Mannltol Sorbitor
+
Trehalose
Maltose
of cockroaches
Blattellageiinanica 'i'-r i 1 1 1 ・ 1
Fructose
+
l
com-
overnight,
repeatedly
was
Bittenspots filter paper.
container
to remain
allowed
1100
memu
Mannitol
of each
tested atatime on each paper 10 to 15. After being dried, the
compounds
varied
!/
n-Glucosamine
Si, S2,
dilutedwith or suspended in 2rnr pound of water. The dilutions and suspensions were dropped on a filter 15cm in diameter, paper making spots of 2cm in diameter.The num-
rm "
+
examine
was
Manno$e
Galactose Fructose L-Sorbose
to
to the extracts
adults
S3 and
f.
R
-
Rhamnose
of
roaches.
a.
tested under
on
some
,1
en
L-Arabinose
quanti-
smaller
sugars
conditions
the responses
within
6.
feedingwas identical to those used
compounds
1
l -+ liI
il
'
t'-
+
/
-
+ -
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ac IF fu im 1965 ij Vol. 16 No.4 Table 3 Order of unit
B.
weight
of
P. fL
L-Arabinose MaltoseL-Arablnose MaltoseL-Arabinose
The
SorbitolSUCTOseFructose Maltose SucroseSerbltel i:1 /
the degree
where
i
i
i Ga]actose
Calactose
by the graded
of
Mannitol
response
number
be
can
plus and
of
estimated
rninus
signs,
bitingresponses of the roaches differedgreatly for different compounds, and that L-arabinose, galactose,rnannitol, sorbitol, maltose, fructose, sucrose and The
table shows
starch
that the
table also
the three species
arnong
disparities of the such
The
effective.
were
that
there
to several
responses
galactose,rnannitol
as mannose,
shows
are
tested
were
114 and 2, and weight
experiment,
second
The
their order
results
each
on
of
are
and
releaser
not
in
soluble
in
n-
is
methanol,
make
oriented
souree
or e!icit
(1962)have
Fraenkel
in the problem
fbr a unit in Table 3.
IV. DIscussloN
as
olfactory
of
of
basisof
or
oligophagous
of
plants under
suLstances
which
act
general nutrients, Fraenkel and Blewett
and
other
however,have (1943),
that the taste and
estimated
and
gustatory'
as
(4)sugars
and
act'
which
(2)water-soluble
attractants,
substances (3)fat-soiuble
already
fbod
classified
the chemical
mono-
green part
feedingstimulants,
to
the
towards
fourheadings; (1)volatile
in Table
to
them
feedingaction.
their
food specificity
even
cause
which
movements
and
words,
insects respond
substances
insectsfeedingon
shown
is shown
typically omnivorous
some
the
raM-
effectiveness
species
these
worlcs
substances
of 1!100, li'10, dilutions
the
with
112 M.
is
soluble
in rice bran, In ether
stimulants
Nayer
stimulants
seven
S3,which but
ether
for feeding of the three species, but it shows no attractive effect (Figs. 2, 3, 4, 5 of the stimulants of this and 7). Some effective type are carbohydrates (Table1, 2 and 3). effeet These results, includingthe attractive D, clearly indicate that of the steam-distillate differentattractants and feeding there are some
some
nose.
In the
and
stimulative
I Sorbitol Sucrose
Mannitol
one.
extract
hexane
Fructose
Fructose
species,
that of neutral
Galactose
)(onnitol
two
other
substances.
a.
g.lR
however, the attractive activity as well as the feeding stimulant one seerns fracto be due to both of the acidic and neutral tions. In these species the feedingstimulant effect of acidic fractien was much greater than
for a
effectiveness
seven
(261)
smell
of
food is probably of great importance even in stered productsinsects which are omnivorous. Murray (1960)has claimed that 7'enebn'o "iolito4 a species of stored products insects, in itslarvalbehaviourpatternof feeding,is to be regarded as not essentially different from other insects, though he has phytophageus given no the
Detheiret
have (1960)
aL
nation
of
which
they elicit
in terms
chemicals
of
the
desig-
responses
from in$ects,They have listed
terms of chemical
as standard
the
made
stimuli
; arrestant,
stimulant
feeding, ovipositional), (rocornotor,
attractant,
repellent
here
obtained
deterrent.The results satisfactorily be discussed
and
may
these terms,
with
in the results, the n-hexane soluble fraction of rice bran (Si) has two effeets, attractant effect and feedingstimulant one, for the cockroaches (Fig.6). Since the residue Ri is As
not rice
the
shown
(Fig.2),
attractive
bran must be extract Si. In
the
adults
the
attractive
effect
by the constituents
caused
indicationas to the chemical nature of the stituents in the food which acts as releaser stances for the initiationand continuation It is chemical
of
be
old
nymphs
of
B.
.uer-
the attractive effect of Si is due to the fraction, and the feedingstimulant effect is attributable to both of the neutral and acidie
stances eding
and
old
nymphs
of the
in the basisof
of extracts
neutrai
adults
of
from the
results
omnivorous' the
of
Si
which stimulant
most (3),
the sugars
components and
the
of
attractant
effect
be due to some subbelongto the heading(1), the feeffect of Si may be due to those
and
D
here,
food can headings as Nayer
non-specificity
the same
under
obtained cockroaches
Fraenkelhave listed.The
7nanica
In the fractiens.
clear
classified
and and
also
of
sub-
feeding.
normal
that
con-
relatea
may
of
S3 may
compeunds
be of (2),and
belongto (4).
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(262 )
衛
There is the lack of specificity eding
stimulants
such
as maltose
fe。 Larabinose
of
,
some
lack
sp
,
of other
the cockroaches
among
substances
,however
mannitol
releaser
chemicals
,
such
are
as
somewhat
galac−
or
−specific at Ieast in the three spe .
highly species cies Table 3) and there are dif6erent responses ( . to the acidic fraction of the n hexane soluble extract of rice bran between B . ge nanica and the other two species . There are also different ,
“
7 lca
〃
A between the newly fraction
adults
’
nymphs hatched
specific repe11ents causal
and
,
/la
1: }Dethier V. G . Brown , L. B . and , ,
C . N . Smlth,
in terms of the responses they elicit fro皿 insects, J. econ . Ent . , 53 : 134 −136 .− 2 )Fraenkel , G . and M . Blewett , ・ (1943) The natural foods and the food require :
chemicals
:
several species of stored products insects , Trans. R . ent . S c. Lond . ,93 457 −490 .− 3 )Mur − ray ,1). 1960 ) The stlmulus to feeding in R . P 〔 ・ ’ , Iarvae f Tenebrio nzo /itot L . J, Ins Physiol . . ,4 − 80 91 .− 4 )Nayer , J. K . and Fraenkel G .任962 } The chemical basls of hostl1ant selection in the silk ・ − worm tb . Bo J.工 ns . Phys め 】 . , yx zori (L . ) , ,8 : 505 ・ 525. ments
of oligo − or monophagous feature
1960 ) The designation of (
deterrents may
,
or
LiTERATURE
in P .amen cana . the species −specificity or the age ・ So specificity of each releaser chem 三 cal which is involved in food se 】ection is not a special feature of oligo ・ insects . phagous or monophagous The restricted distribution of specific attractants and feeding as weH as the wide distribution of stimulants and
galactose and
of
are somewhat
,
to the extract SI and to the acidic
responses
releaser substances
them .
mamito1 how ・ −speci 丘c ln highly species the three species , and there are different re ・ sponses to the acidic fraction A of the n . hexane soluble ger − extract of rice bran between B . inanica and the other two species . There are also different responses to the n ・ hexane extract Sl and to the acidic fraction A between the ’ newly hatched nymphs and adults in P . an ze ・
ever
.
The effects of some tose and
The effects
of n −hexane soluble ificity
among
,
of specificity
Vol .16 No .4
1965 年
物
fructose and sllcrose among the three ies ( Table 1). that there is the It is likely
sorbitol sp
動
生
of
[)
:
:
(,
be the insects .
: :
,
〕
〃
AcKNowLEDGEMENT
〃
The au 山 r wishes to express his hearty thanks to Dr . Syozim Asahina , Department of Medical
Entomo 】gy , National Institute of Health Tokyo , , for his interest and encouragement .
要
. 雑 食性
ゴ キ
旨
3 種 の ,食 物 発 見 と摂 食 反 応 を 引 きお 刺激 の 物 質的 基 礎 は , (1 ) n 一 揮 発 性 の olfactory attractants , (2 ) n .
ブ
リ
こ す 米 ぬ か と. 炭 水化 物 の
SUMMARY
.
キ サ ン
可溶 で キ サ ン 不 溶 , メ タ ール 又 は 水 に 可 溶 の 1ants n ・ キ サ ン 可溶 の ,(3 ) 水 に 不 溶 , ヘ
ヘ
In order to investigate the behaviour pat ・ tern of feedingof three species of cockroaches some iminary experiments were carried out prel. on their responses to olfactory and gustatory of fbod origin . stimuli of the three species to the The responses of rice bran and some carbohydrates constituents indicatethat the chemical basisof their clearly food pre艶 rence can be classified under four headings; (1)volatile substances which are sDl − and act as olfactory attractants uble in n −hexane − −soluble and ( methanol or water 3)n −hexane− 2) ( . substances which act as feeding stimu sDluble and 4) sugars and related compounds lants , ( as stimulants also acting . gustatory feeding of specificity of some fe. There is the lack ,
,
eding
stimulants
sorbitol species
such as maltose
,
Larabinose ,
fructose and sucrose among the three , is likely that there is the . It lack of
ヘ
stimu ・ feeding feeding stimu .
lants ,(4 ) 糖 及 び そ の 類 縁 化 合物 の 刺 激 の 4 項 目 に 分 る. (2 )の 相 当部 分 は (4 )に 属 す る もの で あ ろ う. n ・ キ サ ン 米 ぬ か の ン に は 抽 出物 の 中性 フ ラ ク シ 3 種 と も誘 引 さ れ る . . ル トー.,L ・ ア ラ ビ ノ ース , ソ ース 等 の . ル ビ トール , フ ラ ク トース ,ス ク 炭 水化 物 に し て 3 対 種 と も摂 食 反 応 を 起 す . しか し ガ ラ ク トー ス は , 3 種 の う ち ク ロ ゴ キ ブ リ だ け に 強 く作 用 し , ン トール に は ワ モ ン ゴ キ ブ リ と バ ゴ チ ャ ネ キ ブ リ だ け が よ く反 応 す る . 又 米 ぬ か の n 一 キ サ ン抽 出 物 の 酸 性 フ ラ ク シ ン に対 して チ バ ネ .ゴ . キ ブ リ成 虫 は 誘 引 さ れ 難 い が 他 の 2 種 の 成. 虫 は 誘引 さ れ る. 一 方 , ワ モ ン ゴ キ ブ リ の ふ 化 直 後 の 幼 虫 は n 一 キ サ . ン 抽 出 物 に 誘 引 され る が , そ れ に 対す る 摂. 食反 応 は 弱 く ,そ の 酸 性 フ ラ ク シ ン に は 誘 引 され な い . し か し 成 虫 は い ず れ に も誘 引,摂 食 . 反 応 を 示 し ,特 に 摂 . 食反 応 は ン に対 して 強 か 酸性 フ ラ ク シ た .こ の よ う な age の 類 出来
ヘ
ョ
マ
ス
ロ
,
,
マ
ニ
ヘ
ョ
ャ
,
ヘ
ョ
つ
ョ
差 に よ る反 応 の 相 異 が他の 査 で あ る
2種
で
も あ る か ど うか
.
一
は
未調
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