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The effect of limited milk feeding on the longevity of the house fly, musca domestica L.
Exp. Geront. Vol. 6, pp. 205-210. Pergamon Press1971, Printed in Great Britain
THE EFFECT OF LIMITED LONGEVITY OF THE HOUSE
MILK FEEDING ON THE FLY, MUSCA DOMESTICA
L.
F. H. GRAY*and P. A. BERBERIAN Department of Physiology, Universityof Miami School of Medicine, Miami, Florida 33152, U.S.A.
(Received 22 December 1970) INTRODUCTION IN ORDERto understand aging in the house fly, Musca domestica L., we need to know some of the parameters determining its longevity. Earlier studies to determine the effect of diet on the lifespan of flies showed that milk makes the adult female live longer but has no effect on the lifespan of the adult male (Rockstein, 1957; Rockstein and Lieberman, 1959). Ascher and Levinson (1956) reported that the mean longevity of flies fed milk for two days was similar to that found by Greenberg (1955) who fed milk the entire lifetime of the adults. The present experiment was begun to see whether the longevity of house flies fed milk for three days only at 12 hr following adult emergence was the same as that of flies fed milk constantly throughout adult life.
MATERIALS AND M E T H O D S Adult house flies of the N A I D M strain, long-inbred and reared in this laboratory (from a stock originally supplied by the United States Public Health Service, Savannah laboratory) were maintained at 26.6 + 2°C and 50 per cent R. H., under conditions of continuous lighting. The flies were kept in bronze screen cages (inside dimensions: 7.5 in. per side) with approximately 180 to 200 flies of each sex per cage. The flies were fed ad libitum on sucrose with a constant supply of water. Those flies fed only sucrose and water will be referred to as "sugar flies." Another group of flies had continuous access to Klim (Borden's powdered bovine milk) in addition to the sucrose and water. These flies will be referred to as "constant milk flies". A third group of flies had, in addition to the sucrose and water, access to Klim for three clays only, beginning 12 hours after adult emergence. These flies will be referred to as "3-day milk flies." Dead flies were removed from each cage every 24 hr and sexed, counted and recorded. Mortality curves were constructed from the data and 50 per cent mortalities determined for a total population of fourteen thousand flies. RESULTS
Figure 1 shows that milk feeding has no effect on the longevity of the adult male fly. Figure 2 shows that the 3-day milk female flies have the same increase in longevity as * Present Address: Department of Entomology, Ohio State University, Columbus, Ohio 43212, U.S.A. 205
206
F.H. GRAY AND P. A. BERBERIAN
do constant milk female flies. Tables 1 and 2 summarize the 50 per cent mortalities for each sex on the various dietary regimens.
loo
90
/
80
./
@
70
6O o eCM
o
E
5o
o3DM
c o
a.
uCS 40
30
20
10
/
1 l l 12
6
l
i
l
18
Age,
l 24
l
l l 30
l
[ 36
l
l l 42
l 48
54
day
FIG. 1. T h e effect of diet on the longevity of male house flies.
T A B L E 1.
M O R T A L I T Y DATA OF MALE HOUSE FLIES O N VARIOUS DIETARY R E G I M E N S : CONSTANT M I L K ,
SUGAR AND W A T E R
(CM);
THREE DAYS OF M I L K W I T H
CONSTANT SUGAR AND W A T E R
(3 D M ) ;
CONSTANT SUGAR AND W A T E R ( C S )
Diet
Cages
N u m b e r of flies
50 per cent mortality
Maximum longevity
CM
15
3063
11"5
36
3DM
15
2769
11"8
33
7
1286
11"5
24
CS
207
T H E EFFECT OF LIMITED M I L K FEEDING
lOG
9C
/
80
70
60
2 :s E E Q e,
50
• CM o 3DM n CS
40
30
20
I0
S
12
18
24
30 Age,
F I G . 2.
The effect of diet on
the
36
42
4.8
54
60
66
day
longevity of female house flies.
T A B L E 2. M O R T A L I T Y DATA OF FEMALE HOUSE FLIES ON VARIOUS DIETARY REGIMENS: CONSTANT MILK, SUGAR AND WATER ( C M ) ; THREE DAYS OF M I L K W I T H CONSTANT SUGAR AND WATER (3 DM); CONSTANT SUGAR AND WATER ( C S )
Diet
Cages
Number of flies
CM 3DM CS
15 15 7
2981 2706 1180
50 per cent mortality 21-0 23.1 15-8
Maximum longevity 64 52 36
DISCUSSION T h e results of the present experiment are in agreement with those of Rockstein (1957) and Rockstein and Lieberman (1959) who showed that milk increases the longevity of the female house fly and has no effect on that of the male. T h e 50 per cent mortalities for males and females on constant milk diets are in agreement with those reported by Bhatnagar, Rockstein and Dauer (1965).
208
F . H . GRAY AND P. A. BERBERIAN
In this experiment we have shown that feeding milk for three days has the same beneficial effects on the longevity of the female as does constant milk. In understanding this result it is important to realize that female flies (1), need milk to develop eggs (personal observations), (2) have a fully mature batch of eggs by the fourth day of postemergence when on a milk diet, (3) retain eggs unless a suitable oviposition site is provided (Morrison and Davies, 1964; and personal observations), (4) were not provided with a suitable environment for laying eggs in the present experiment. It should also be noted that Strangeways-Dixon (1961) and E. T h o m s o n and Moller (1963) showed that Calliphora erythrocephala females with ripe eggs do not ingest protein food but just sugar and water. W e suspect that females on either constant milk or 3-day milk diets ingest enough milk shortly after emergence for the maturation of a batch of eggs, because of the lack of an oviposition site do not lay their eggs, and do not drink more milk but only sugar and water. Thus, access to milk after the first three to four days of post-emergence has no additional effect since it is not drunk and cannot be utilized. T h e effect of increased longevity is established within the first three days after 12 hr post-emergence and explains why constant milk and 3-day milk females have the same increased longevities. It was observed that a minimal amount of egg laying did take place on the gauze around the ends of the water vials used to supply water to the flies, and this may explain the differences in the average maximum longevities of the females on constant milk and 3-day milk. We postulate that those females on constant milk which did lay eggs were able to drink more milk, develop another batch of eggs and experience another increase in their longevity. T h o s e females on a 3-day milk diet which laid eggs did not have further access to milk and thus could not ingest more milk and benefit in the same way. Experiments have been initiated in this laboratory to test the effect of egg laying on the female's longevity (to be published). Acknowledgements--This investigation was supported in part by funds from Research Grant HD-00571 and Training Grant HD-00142 from the National Institute of Child Health and Human Development of the United States Public Health Service. We are indebted to Dr. Morris Rockstein for making available the materials and facilities utilized in this experiment, and we especially thank him for his comments and interest. REFERENCES
ASCHER,K. R. S. and LEVINSON,Z. H. (1956)Rivista Di Parassitologia. X V I I - N 4 , 217. BHATNAQAR,P. L., ROCKSTEIN,M. and DA~a, M. (1965). Exp. Gerontol. 1, 149. GREENBERG, G. (1955)ft. Econ. Ent. 48, 125. MoamsoN, P. E. and DAVIES, D. M. (1964) Nature 201, 194. ROCKSTErN, M. (1957)ft. Gerontol. 12, 153. ROCKSTEIN, M. and LmBm~WAN,H. M. (1959) Musca domestica L Gerontologia 3, 23. STP,ANGEWAYS,D. J. (1961) J. Exp. Biol. 38, 225. THOMSON, E. and MOLLER, I. (1963) J. Exp. Biol. 41, 301. S u m m a r y ~ F e m a l e house flies live as long on a 3-day milk supplement beginning 12 hr after emergence as they do on a constant milk supplement throughout their entire life span. Male house flies are not affected. The beneficial effects on female house fly longevity are indicative that utilization of milk is intimately related to egg development, and this in turn related to increased longevity.
T H E EFFECT OF L I M I T E D M I L K FEEDING
Female house flies on either constant or 3-day milk regimens ingest enough milk shortly after emergence for the maturation of a batch of eggs, and because of the lack of suitable oviposition site do not lay their eggs. Rather, they harbor a fully developed batch of eggs, and do not ingest more milk. Since milk is not ingested and utilized after the first three to four days of postemergence, the effect of increased longevity due to milk supplementation is established within this time via egg development. This explains the similar increased longevities of females on constant and 3-day milk dietary regimens. R d s u m ~ Les mouches domestiques femelles auxquelles u n suppldment de lait est servi pendant 3 jours /~ partir de la 12e heure suivant leur dclosion, vivent aussi longtemps que leurs congdnbres qui re~oivent u n suppldment de lait constant tout au long de leur vie. Cet effet ne s'observe pas chez les mouches domestiques males. L'effet bdndfique sur la longdvit6 de la mouche domestique femelle indique que l'assimilation du lait est en corrdlation dtroite avec le ddveloppement des oeufs, ce qui, ~ son tour, est en corrdlation avec une longdvitd accrue. Qu'elles soient soumises au rdgime lactd de 3 jours ou au rdgime constant, les mouches domestiques femelles consomment, peu de temps apr~s leur dclosion, suffisamment de lair pour pourvoir A la maturation d ' u n lot d'oeufs, mais ne pendent pas ceux-c~, faute d ' u n site d'oviposition approprid. Elles hdbergent plut6t u n lot d'oeufs complAtement ddveloppd et ne consomment plus de lair. Le lair n'dtant plus consommd et utilisd aprAs les trois ou quatre premiers jours suivant l'dclosion, l'effet de longdvit6 accrue dO au suppldment de lait est 6tabli dans ce laps de temps via le ddveloppement des oeufs. Ceci explique l'augmentation similaire de la longdvitd chez les femelles soumises au rdgime lactd constant et chez celles soumises au rdgime lactd de 3 jours. Z u s a m m e n f a s s u n g - - W e i b l i c h e Hausfliegen, welche 12 Stunden nach dem Schlfipfen ffir 3 Tage Milchzusatz zur Nahrung bekommen, leben ebenso lange wie solche, die Milchzusatz wghrend des gesamten Lebens erhalten. M~mliche Hausfliegen sind nicht betroffen. Die positive Wirkung auf die Lebensdauer der weiblichen Hausfliegen deutet auf eine enge Beziehung des Milchverbrauchs zur Eientwicklung, u n d diese wiederum auf eine Beziehung zur erh6hten Lebensdauer. Weibliche Hausfliegen mit konstanter oder 3-tiigiger Milchnahrung nehmen kurz nach dem Schlfipfen genug Milch zur Reifung vieler Eier auf, welehe sie wegen Fehlens einer geeigneten Legestelle jedoch nicht legen. Sie bewahren daher die vollentwickelten Eier auf u n d nehmen keine weitere Milch zu sich. Da nach den ersten 3-4 Tagen nach dem Schliipfen keine Milch mehr aufgenommen u n d gebraucht wird, wird die Wirkung auf die erh6hte Lebensdauer durch Milchzusatz w~ihrend dieser Zeit fiber die Eientwicklung entfaltet. Dies erkl/irt die bei Weibchen mit konstanter oder 3-t/igiger Milchnahrung in gleicher Weise erh6hte Lebensdauer. Pe31oM~, CaMKH KOMHaTHO~ MyXH MOFyT XHTb C 3-]~HeBHI,IMPI~O6aBKaMH MOHOKa, HasHHa~ C 12 qacoB n e e . h e BM.r[eTa, TaK)Ke ]~oJ1ro, KaK H C HOC'rO~HH~MH ~ o 6 a m z a ~ MOHOKa a
IIpo]IoH)KeHHe Bcero cBoero ~K3HeHHOFO Ip~IKHa. C ~)yro~ CTOpORM, KOpM.rlem4e M O H O K O M He OKa3MBaeT B.wH4,qHHHHa H p o ~ o ~ . F ~ H O C T b ~ [ 3 H H CaMI~OB KOMHaTHOI~ MyXH. ]~HaFOTBOpHO¢ BJERflHHe aH ~iHTeHbHOCTb ~[~3HH CaMOK KOMHaTHOI~ MyXH CBI~eTeH~TByOT O TOM, HTO HCFIOHb3OB~I4e MOJIOKa TeCHO CBff3aHO C pa3BHTHeM ~IHI~, a IIOCJIe]]~H~, B CBOIO o~Iepe]Ib, OKa3MBaeT ~ e Ha y ~ ' m H e H H e npo~oJLxorre~bHOCTH )KH3HH. CaMKH KOMHaTHO~ MyXH Ha HOCTORHHOi~HYm 3-]IHeBHOI~ MOJIO~IHOI~~HeTe n o r n o I ~ a ~ o T ]IOCTaTOv~HO MOHOKa B c ~ o p e HoCHe B]~HeTa, ~rro6M o 6 ¢ c n e m ~ T b co3pCBaHHe napTHH smal; o ] l H a g o IIpH OTCyTCTBI4H Ho]~xo]IRIlIero MCCTa ] U ~ OTKHag~KHOHH ~ He OTgHa]Ibma~OT, HO c o x p a H ~ O T HapTHIO BrloHHe pa3BHTI,[X ~ H He IIOFHOLI~aIOT6on.me MOHOKa.
209
210
F. H . GRAY AND P. A. BERBERIAN I'IocKOHBKy MOIIOK0 HO IlOFJI0ma~c~I H He HCn07tb3yeTCJl HOCHe gCpBblX T ~ X H~H •leTbIpOX ~ I O ~ HOG/IC Bbl/IeTa, ~e~CTBHe y~/IHHeHHO~ IlpO,~O.rDKHTebHOCTH ~ I 3 H H , B/d3BaHHOC ~O6aBKaMH MOHOKa, BbIHBJLqOTCflB TOtIOHH¢ 3TOFO cpoxa IIyTCM pa3BHTH.q flh~. ~THM O6'I~RCHJICTC~I O~HHaKOBMC y~nHHOHH,q HpoRoJI~,HTC0"II, HOCTH ~KH3HH CaMOK KaK n p H HOCTOflHHOM, TaX H IlpH 3-~HeBHOM ,~HOTH~IeCKHXyCYlOBH,qX.
THE EFFECT OF LIMITED LONGEVITY OF THE HOUSE
MILK FEEDING ON THE FLY, MUSCA DOMESTICA
L.
F. H. GRAY*and P. A. BERBERIAN Department of Physiology, Universityof Miami School of Medicine, Miami, Florida 33152, U.S.A.
(Received 22 December 1970) INTRODUCTION IN ORDERto understand aging in the house fly, Musca domestica L., we need to know some of the parameters determining its longevity. Earlier studies to determine the effect of diet on the lifespan of flies showed that milk makes the adult female live longer but has no effect on the lifespan of the adult male (Rockstein, 1957; Rockstein and Lieberman, 1959). Ascher and Levinson (1956) reported that the mean longevity of flies fed milk for two days was similar to that found by Greenberg (1955) who fed milk the entire lifetime of the adults. The present experiment was begun to see whether the longevity of house flies fed milk for three days only at 12 hr following adult emergence was the same as that of flies fed milk constantly throughout adult life.
MATERIALS AND M E T H O D S Adult house flies of the N A I D M strain, long-inbred and reared in this laboratory (from a stock originally supplied by the United States Public Health Service, Savannah laboratory) were maintained at 26.6 + 2°C and 50 per cent R. H., under conditions of continuous lighting. The flies were kept in bronze screen cages (inside dimensions: 7.5 in. per side) with approximately 180 to 200 flies of each sex per cage. The flies were fed ad libitum on sucrose with a constant supply of water. Those flies fed only sucrose and water will be referred to as "sugar flies." Another group of flies had continuous access to Klim (Borden's powdered bovine milk) in addition to the sucrose and water. These flies will be referred to as "constant milk flies". A third group of flies had, in addition to the sucrose and water, access to Klim for three clays only, beginning 12 hours after adult emergence. These flies will be referred to as "3-day milk flies." Dead flies were removed from each cage every 24 hr and sexed, counted and recorded. Mortality curves were constructed from the data and 50 per cent mortalities determined for a total population of fourteen thousand flies. RESULTS
Figure 1 shows that milk feeding has no effect on the longevity of the adult male fly. Figure 2 shows that the 3-day milk female flies have the same increase in longevity as * Present Address: Department of Entomology, Ohio State University, Columbus, Ohio 43212, U.S.A. 205
206
F.H. GRAY AND P. A. BERBERIAN
do constant milk female flies. Tables 1 and 2 summarize the 50 per cent mortalities for each sex on the various dietary regimens.
loo
90
/
80
./
@
70
6O o eCM
o
E
5o
o3DM
c o
a.
uCS 40
30
20
10
/
1 l l 12
6
l
i
l
18
Age,
l 24
l
l l 30
l
[ 36
l
l l 42
l 48
54
day
FIG. 1. T h e effect of diet on the longevity of male house flies.
T A B L E 1.
M O R T A L I T Y DATA OF MALE HOUSE FLIES O N VARIOUS DIETARY R E G I M E N S : CONSTANT M I L K ,
SUGAR AND W A T E R
(CM);
THREE DAYS OF M I L K W I T H
CONSTANT SUGAR AND W A T E R
(3 D M ) ;
CONSTANT SUGAR AND W A T E R ( C S )
Diet
Cages
N u m b e r of flies
50 per cent mortality
Maximum longevity
CM
15
3063
11"5
36
3DM
15
2769
11"8
33
7
1286
11"5
24
CS
207
T H E EFFECT OF LIMITED M I L K FEEDING
lOG
9C
/
80
70
60
2 :s E E Q e,
50
• CM o 3DM n CS
40
30
20
I0
S
12
18
24
30 Age,
F I G . 2.
The effect of diet on
the
36
42
4.8
54
60
66
day
longevity of female house flies.
T A B L E 2. M O R T A L I T Y DATA OF FEMALE HOUSE FLIES ON VARIOUS DIETARY REGIMENS: CONSTANT MILK, SUGAR AND WATER ( C M ) ; THREE DAYS OF M I L K W I T H CONSTANT SUGAR AND WATER (3 DM); CONSTANT SUGAR AND WATER ( C S )
Diet
Cages
Number of flies
CM 3DM CS
15 15 7
2981 2706 1180
50 per cent mortality 21-0 23.1 15-8
Maximum longevity 64 52 36
DISCUSSION T h e results of the present experiment are in agreement with those of Rockstein (1957) and Rockstein and Lieberman (1959) who showed that milk increases the longevity of the female house fly and has no effect on that of the male. T h e 50 per cent mortalities for males and females on constant milk diets are in agreement with those reported by Bhatnagar, Rockstein and Dauer (1965).
208
F . H . GRAY AND P. A. BERBERIAN
In this experiment we have shown that feeding milk for three days has the same beneficial effects on the longevity of the female as does constant milk. In understanding this result it is important to realize that female flies (1), need milk to develop eggs (personal observations), (2) have a fully mature batch of eggs by the fourth day of postemergence when on a milk diet, (3) retain eggs unless a suitable oviposition site is provided (Morrison and Davies, 1964; and personal observations), (4) were not provided with a suitable environment for laying eggs in the present experiment. It should also be noted that Strangeways-Dixon (1961) and E. T h o m s o n and Moller (1963) showed that Calliphora erythrocephala females with ripe eggs do not ingest protein food but just sugar and water. W e suspect that females on either constant milk or 3-day milk diets ingest enough milk shortly after emergence for the maturation of a batch of eggs, because of the lack of an oviposition site do not lay their eggs, and do not drink more milk but only sugar and water. Thus, access to milk after the first three to four days of post-emergence has no additional effect since it is not drunk and cannot be utilized. T h e effect of increased longevity is established within the first three days after 12 hr post-emergence and explains why constant milk and 3-day milk females have the same increased longevities. It was observed that a minimal amount of egg laying did take place on the gauze around the ends of the water vials used to supply water to the flies, and this may explain the differences in the average maximum longevities of the females on constant milk and 3-day milk. We postulate that those females on constant milk which did lay eggs were able to drink more milk, develop another batch of eggs and experience another increase in their longevity. T h o s e females on a 3-day milk diet which laid eggs did not have further access to milk and thus could not ingest more milk and benefit in the same way. Experiments have been initiated in this laboratory to test the effect of egg laying on the female's longevity (to be published). Acknowledgements--This investigation was supported in part by funds from Research Grant HD-00571 and Training Grant HD-00142 from the National Institute of Child Health and Human Development of the United States Public Health Service. We are indebted to Dr. Morris Rockstein for making available the materials and facilities utilized in this experiment, and we especially thank him for his comments and interest. REFERENCES
ASCHER,K. R. S. and LEVINSON,Z. H. (1956)Rivista Di Parassitologia. X V I I - N 4 , 217. BHATNAQAR,P. L., ROCKSTEIN,M. and DA~a, M. (1965). Exp. Gerontol. 1, 149. GREENBERG, G. (1955)ft. Econ. Ent. 48, 125. MoamsoN, P. E. and DAVIES, D. M. (1964) Nature 201, 194. ROCKSTErN, M. (1957)ft. Gerontol. 12, 153. ROCKSTEIN, M. and LmBm~WAN,H. M. (1959) Musca domestica L Gerontologia 3, 23. STP,ANGEWAYS,D. J. (1961) J. Exp. Biol. 38, 225. THOMSON, E. and MOLLER, I. (1963) J. Exp. Biol. 41, 301. S u m m a r y ~ F e m a l e house flies live as long on a 3-day milk supplement beginning 12 hr after emergence as they do on a constant milk supplement throughout their entire life span. Male house flies are not affected. The beneficial effects on female house fly longevity are indicative that utilization of milk is intimately related to egg development, and this in turn related to increased longevity.
T H E EFFECT OF L I M I T E D M I L K FEEDING
Female house flies on either constant or 3-day milk regimens ingest enough milk shortly after emergence for the maturation of a batch of eggs, and because of the lack of suitable oviposition site do not lay their eggs. Rather, they harbor a fully developed batch of eggs, and do not ingest more milk. Since milk is not ingested and utilized after the first three to four days of postemergence, the effect of increased longevity due to milk supplementation is established within this time via egg development. This explains the similar increased longevities of females on constant and 3-day milk dietary regimens. R d s u m ~ Les mouches domestiques femelles auxquelles u n suppldment de lait est servi pendant 3 jours /~ partir de la 12e heure suivant leur dclosion, vivent aussi longtemps que leurs congdnbres qui re~oivent u n suppldment de lait constant tout au long de leur vie. Cet effet ne s'observe pas chez les mouches domestiques males. L'effet bdndfique sur la longdvit6 de la mouche domestique femelle indique que l'assimilation du lait est en corrdlation dtroite avec le ddveloppement des oeufs, ce qui, ~ son tour, est en corrdlation avec une longdvitd accrue. Qu'elles soient soumises au rdgime lactd de 3 jours ou au rdgime constant, les mouches domestiques femelles consomment, peu de temps apr~s leur dclosion, suffisamment de lair pour pourvoir A la maturation d ' u n lot d'oeufs, mais ne pendent pas ceux-c~, faute d ' u n site d'oviposition approprid. Elles hdbergent plut6t u n lot d'oeufs complAtement ddveloppd et ne consomment plus de lair. Le lair n'dtant plus consommd et utilisd aprAs les trois ou quatre premiers jours suivant l'dclosion, l'effet de longdvit6 accrue dO au suppldment de lait est 6tabli dans ce laps de temps via le ddveloppement des oeufs. Ceci explique l'augmentation similaire de la longdvitd chez les femelles soumises au rdgime lactd constant et chez celles soumises au rdgime lactd de 3 jours. Z u s a m m e n f a s s u n g - - W e i b l i c h e Hausfliegen, welche 12 Stunden nach dem Schlfipfen ffir 3 Tage Milchzusatz zur Nahrung bekommen, leben ebenso lange wie solche, die Milchzusatz wghrend des gesamten Lebens erhalten. M~mliche Hausfliegen sind nicht betroffen. Die positive Wirkung auf die Lebensdauer der weiblichen Hausfliegen deutet auf eine enge Beziehung des Milchverbrauchs zur Eientwicklung, u n d diese wiederum auf eine Beziehung zur erh6hten Lebensdauer. Weibliche Hausfliegen mit konstanter oder 3-tiigiger Milchnahrung nehmen kurz nach dem Schlfipfen genug Milch zur Reifung vieler Eier auf, welehe sie wegen Fehlens einer geeigneten Legestelle jedoch nicht legen. Sie bewahren daher die vollentwickelten Eier auf u n d nehmen keine weitere Milch zu sich. Da nach den ersten 3-4 Tagen nach dem Schliipfen keine Milch mehr aufgenommen u n d gebraucht wird, wird die Wirkung auf die erh6hte Lebensdauer durch Milchzusatz w~ihrend dieser Zeit fiber die Eientwicklung entfaltet. Dies erkl/irt die bei Weibchen mit konstanter oder 3-t/igiger Milchnahrung in gleicher Weise erh6hte Lebensdauer. Pe31oM~, CaMKH KOMHaTHO~ MyXH MOFyT XHTb C 3-]~HeBHI,IMPI~O6aBKaMH MOHOKa, HasHHa~ C 12 qacoB n e e . h e BM.r[eTa, TaK)Ke ]~oJ1ro, KaK H C HOC'rO~HH~MH ~ o 6 a m z a ~ MOHOKa a
IIpo]IoH)KeHHe Bcero cBoero ~K3HeHHOFO Ip~IKHa. C ~)yro~ CTOpORM, KOpM.rlem4e M O H O K O M He OKa3MBaeT B.wH4,qHHHHa H p o ~ o ~ . F ~ H O C T b ~ [ 3 H H CaMI~OB KOMHaTHOI~ MyXH. ]~HaFOTBOpHO¢ BJERflHHe aH ~iHTeHbHOCTb ~[~3HH CaMOK KOMHaTHOI~ MyXH CBI~eTeH~TByOT O TOM, HTO HCFIOHb3OB~I4e MOJIOKa TeCHO CBff3aHO C pa3BHTHeM ~IHI~, a IIOCJIe]]~H~, B CBOIO o~Iepe]Ib, OKa3MBaeT ~ e Ha y ~ ' m H e H H e npo~oJLxorre~bHOCTH )KH3HH. CaMKH KOMHaTHO~ MyXH Ha HOCTORHHOi~HYm 3-]IHeBHOI~ MOJIO~IHOI~~HeTe n o r n o I ~ a ~ o T ]IOCTaTOv~HO MOHOKa B c ~ o p e HoCHe B]~HeTa, ~rro6M o 6 ¢ c n e m ~ T b co3pCBaHHe napTHH smal; o ] l H a g o IIpH OTCyTCTBI4H Ho]~xo]IRIlIero MCCTa ] U ~ OTKHag~KHOHH ~ He OTgHa]Ibma~OT, HO c o x p a H ~ O T HapTHIO BrloHHe pa3BHTI,[X ~ H He IIOFHOLI~aIOT6on.me MOHOKa.
209
210
F. H . GRAY AND P. A. BERBERIAN I'IocKOHBKy MOIIOK0 HO IlOFJI0ma~c~I H He HCn07tb3yeTCJl HOCHe gCpBblX T ~ X H~H •leTbIpOX ~ I O ~ HOG/IC Bbl/IeTa, ~e~CTBHe y~/IHHeHHO~ IlpO,~O.rDKHTebHOCTH ~ I 3 H H , B/d3BaHHOC ~O6aBKaMH MOHOKa, BbIHBJLqOTCflB TOtIOHH¢ 3TOFO cpoxa IIyTCM pa3BHTH.q flh~. ~THM O6'I~RCHJICTC~I O~HHaKOBMC y~nHHOHH,q HpoRoJI~,HTC0"II, HOCTH ~KH3HH CaMOK KaK n p H HOCTOflHHOM, TaX H IlpH 3-~HeBHOM ,~HOTH~IeCKHXyCYlOBH,qX.