s al tz Vol. 16 No.4

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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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!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 )



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



fructose   and   sllcrose  among  the three ies ( Table  1).  that  there   is   the   It  is  likely

sorbitol sp

 





  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 .



 

. 雑 食性

   

ゴ キ



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 .





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



   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



 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



































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