JPRS ID: 8624 USSR REPORT BIOMEDICAL AND BEHAVIORAL SCIENCES
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20 AUtiUST i9T9 t FOUO Sl79 ) i OF S
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~ ~Oit O~~ICIA1. US~ ONI.Y
JPRS L/8624
20 August 1979 ~
USSR Re ort
p
BIOMEDICAI AND BEHAVIORAL SC:IENCES
(FOUO 5/79)
FBIS FOREIGN BROADCAST INFORMATION SERVICE
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JPRS L/8624
� 20 August 1.979
USSR REPORT '
BIOM~DICAL A~tD BEHAVIORAI. SCI~NC~S
(FOUO 5/79)
This serial publication contains arCicles, abstracts of articles and newa
itema from USSR scient3fic and technical ~ournals on Che apecific sub~ects
refleceed ia the table of contenta.
Photoduplications of foreign-language sources may be obeained from the
Photaduplication Service, Library of Congreas, Washi:~gton, D. C. 20540.
Requcsts should provide adequate identi�icaeion both as to the source and
the individual arCicle(s) desired.
CONIENTS PAGE
AGROTECHNOLOf3Y
Raising the Heat and Drought Resistance of Spring Wheat
by a Complex Method
- (P. A. Genkel~, et al.; FIZIOLOGIYA RASTE~iTY, No 3,
_ 1979) ~
ENZYMOLO(~Y .
InYluence of Effectors on the Folding of an Iumiobilized -
Protein (Trypsin)
(V. 4. Mozhayev, et al.; MOLII{fJI~ARNAYA BIOIAGIYA,
No 3, 1979) 11
MARINE MfANQ~17,S �
On the Problem of Chemorece~,tion in Dolphin~
(G. B. Agarkov, S. A. Gilevich; `,IESTNIK Z00`LOGII,
No 3, 1979) 23
PLA1~ PHYSIOLOGY
Determination of the Threshold of Wheat Sgrout and Root
Sensitivity to Magnetic Field Inter+nity
(N. I. Bogatina, et al.; FT,ZIOLOGI'!A RASTENIY, No 3,
1979) 35
- a- [IT.I - US5R - 21A S&T FOliO~
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N'utt urr~lt;~.~w Ub~ UNLY
CON'I'EN~T'S (Con~inued) ~~e
SC7.EN~.'ISTS AND SC~ENTZFIt", ORGAN27.,ATIQNB
Seaand A~.~.-Un~.orl Conference on Probleme in Bi,omech~nics
V. Knete; MEKE~A.NSKA KOMPOZITNYKH MATERIALOV,
No 3, ~~979) 42
Renolution o:f the Second A].l-Union Conference on Yr::blems
in Biomecklanics ~
(MEY.~ANIKA xot~OZ7LTNYIffi MATF~tIALOV, No 3, 1979 48
PUB,LICATTONS
Chemica:L Prevention of R~diation Contamination
(A. S. Mozzhukhin, F'. Yu. Rachinakiy; I~IMICF~SKAYA
~xoFU~rr~a Ranr.~~siorrrr~a~ PoRaz~nr, i979 ) . . . . . 50
Chemtcal and Biological Agents for Plant Pro~tection:
A Short Handbook -
(N. V. Sazonova; F~IMIC~{IYE I BIOLO(~IC~SttIYE
SREDSTVA ZASHCHITY RASTENIY~ 1978~ 53
Clinical Methods, Diagnoeis and Treatment for Attacks
by Toxic Chemtcal Agentb
(Yu. N. Stroykov; KLINIKA DIAGNOSTIKA I LEC~NZYE
PORA'LI~1I7C OTRAXI,YAYUSHCffiMI VESHCH~TVAMT, ~978) 58
- b -
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AGROTECHNOL~GY
UDC 581.1.032.3
R~IISING TH~ H~AT AND DROUGHT RESISTANCE OF SpR'[NG WHEAT BX A COMPLEX N~THOD
Moscow F'IZZOLOGIYA RASTEN~Y in Russian No 3, 1979 pp 641-647
[Article by P. A. Genkel', K. A. Badanova, L. D. Prusakova, and K. S.
Bokarev, USSR Academy of Sciences Institute of Plant Physiology imeni K. A.
Timiryazev, Moscowj
(Text] A new method for raising the heat and drought
resistance of cereal cro~s, a combination of presowing
hardening of the seeds and semidry processing with
re~ardants,is proposed.
Presowing drought-hardening of Albidum 43 wheat caryopses
by Genk~l's method coupled with their treatment by
5 percent CCC solutinn or 0.5 percent BES solution
stimulated stem growth and maturation in the presence
of both optimum soil moisture and dry soil. The
combination of harc~ening and retardant processing
caused the tilleri.ng node to develop deeper in the
soil in comparison with control plants and with plants
treated with retardants alone. This was accompanied
by a rise ;n the resistance of leaf cells to dehydration
and high temperature in the tillering and stem extension
phases. D�ry soil decreased the grain yield from un-
hardened plants qrown in a phytotron by 85 percent,
that from hardened plants grown from seeds treated with
CCC by 20 percent, and seeds treated with BES by only
10 percent, which considerably reduced the yield loss
in comparison with using each method for raising
resistance to heat and drought separately.
Raising plant resistance to heat and dehydration continues to be one of the
most important problems of plant physiology.
The resistance of plants to drought--that is, to heat and dehydration--is
raised ~n many regions of the Soviet Union with the help of agrotechnical
techniques, b}~ breeding drought and heat resistant varieties, and by means
1
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of o~her metihod~ for raising plant drought resistance. Thus Genkel' (1)
gugc~aa~ed a m~t~?od for pregowing drough~ hardening of sQeds. 'Chis me~hod,
which relies on the significanti adaptiabili~y o� plantis, concurrently in-
creaseg hea~ and droughti resis~ance.
Th~ insti~utie's laboratory of planti drouqht resiatance has revoaled a number
of ana~omical, morphoingical, and physiological charactieristics of hardened
pl~nrs which permit tihem to endura dry soil and atmosph~re better. Owing ~o
~rotective adaptive reactions, hardened plants main~ain a high level of
synthesis, which in ~urn insureg high metabolism and causes changes in a `
number of properties of protop'lasmic organelles, promoting high productivity
n� hardened plants expnsed to drought (2-4). A mechanized seed har.�dening
process proposed by one o� ~he authors (5) permits broader applicati.ton of
the p1an~ hardening method in the �arming practices employed in arid
regions nf r~ur country.
A new way �or rai.sing plant drough~ resistance a~ppeared with discovery of
re~~rdan~s, chlorocholine chloride in particular (6). As we know, chlorocholine,
chloride (CCC), which has an influence on many physiological processes in
pl~ntis, inhibits stem grow~h and, at tihe same time, reduces the demand of
stems for water and makes them more resistant to wilting (6,7). The ,
resistiance of plants treated with this preparation to drought increases
because they experience activatiion of a number of protective adaptive
reactiaons resulting in development of properties typical of drought
resistiant plants: a well developed, deeply psnetrating root system, low
i~eight, reduced transpiration intensity, increased tissue water content,
and reduced permeability of protoplasm to electrolytes (8-11).
A higher water cont~nt and better water metabolism were observed in plants
='I sprayed with CCC. Good plant water metabolism promotes a rise in the water
retention capacity of plants experiencing ~'ought conditions, more economical
water consumption, and grea~er activity of some key enzymes of respiration
and ~f phosphorus and carbohydrate metabolism--ATP-ase, catalase, and
cy*ochrome oxidase (12,13).
It caii be hypothesized that the drought and heat resistance of plants could ,
be raised even more by combining the physiological method for hardening
caryopses against drouqht (wetting the seeds and subsequently drying them) _
~ with a chemical method (treatinq them with retardants before sowing).
Because CCC has a favorable effect on water metabolism in plants experiencing
c3�:ought, we could suggest the nypothesis that the zetardant's effect on
arought resistance would manifest itself even more in hardened plants.
b1e thods
Zwo retardants were used in the experiment~-- crystalline trimethyl-S-
chlorethylammonium chloride produced by the Kemerovo Nitrogen Fertilizer
Plant, and a preparation synthesized by one of the authors of ti:i.s article
(14) named BES (dimethyl-R-bromethylsulfonium bromide). BES is a crystalline
. 2
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~ subg~ance ~hat dis~olves well in water and is similar in chemical qtructure
~o CCC. I~~ formula is
((Cfiy) SCHaCH~BrJ+ Or`.
~
The prepara~ion demonstr~ted high re~ardant activity agains~ peas and
barley (14,15).
' The research was conducted in 1975-1977. Seeds were subjected to semidry
treatment by ~he method proposed by Zadontaev et al. (16).
The ob3ect of research was Albidum 43 spring wheat. Unhardened caryopses
and caryopses subjected to presowing drought hardening according to Genkel's
method were shaken together with the re~ardants for 6 hours in a revolving
drum. Ten mi~.liliters of 0.5 percent BES solution or 5 percent CCC solution
was added to ever~+ 100 gm caryopsesj caryopses sprayed with the same
quantity of water served as the control. Following treatment, the caryopses
were planted in pots of soil. The experiments were performed in a green-
house following the design shown in Table 1.
Table 1. Experimental Design
~l~ I~QCJItlb~l`BHUN 06Q~601KU CCMAII ~5'
N~ I A E)lIBJNHOCT6 IIOVBW I1pH
sapHaHra ~eKil~~~ L'kl lyp;ItljH8~H1111 ~1dCiCHHfI
!~O I~lHMlAIO CCC b~[,
z -'r - ~ - To ine~ (~,geaia (6)
y -I- - - To ~ce ~7) rpyGxosaHes~ I8)
s -f� -+f- - To ~ce~ (7~en~a (~i)
I 8 -f- - To *e ~83W TPyGK088HNi1 ~g~
!0 _ ~Og~ ece a~enia (~6)
-f- To me (7~
~1 -i 40"~ c ~i~t Tpy6xoeaxN~ (8~
i2 - To ~ce ~ ~ 1
Key: -
l. Variant 6. Throughout
2. Presowing seed treatment 7. As above
3. Hardening, Genkel " s method 8. Beginning with the
4. BES stem extension
5. Soil moisture content during phase
plant growth
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- The 1976 experiment diEfered from the others in ~hati drought condi~ions
ware crea~ed in ~he soil at ~he end o� ~he tiillering phase and a�fected
the beginning of ~he �lowering phase. The plan~s were not watered for 7
days. Soi1 moisture in the pots was dropped to 10 percenti. Then watering
- was resumed. The exper3mental plants were subsequen~ly kept under obser- '
vation, and ~heir yield was determined ati the end o� the vegetative period.
The direct effect soil dryness had on experimental plan~s was studied in
~he homeastatic environment o� a Ahytotron in January-March 1976, using the
same experimental design. The plants wera grown ati 22� during the day and
16� at night. P1ant illumination was 200,000 Qrgs�cm 2�sec-1 for 16 hours
(xenon lamps). In ~he ear formation stags ~he plants were sub3ected
to soil drought (watering was interrupted for 11 days). This time soil
moi~ture dropped to 35 percent of its total moisture capacity, which
corresponded to a soil water concentration of 15 percent. The moisture
content of the soil of control plants was kept at 70 percent during this
time by watering.
All experiments had a fivefold replication. Heat resistance was determined
by Matskov's method (17) while resistance of ce11s to dehydration (18) was
determined by the dessiaator method, the criterion being the time of death
of cells experiencing dehydration in the presence of diluted sulfuric acid
, in the tillering and stem extension phases. Determination of the length nf
the underground internode, which was an indication of the depth of the
tillering node, involved fiftyfold replication.
- Results
Hardened caryopses produced sprouts 2-3 days earlier than did unhardened
plants. The retardants CCC and BES did not alter the sprouting time. Growth
inhibition was observed in the first stages of growth of plants frvm
caryopses treated with CCC and grown in the presence oE 70 percent soil
moisture; growth was stimulated in subsequent stages, and the height of the
plants evened out (see Figure). In contrast to this, the retardant BES
stimulated growth as early as in the first stages of plant growth, and it
continued to have this effect until the end of veqetative growth.
Hardened plants in all experimental variants entered the tillering and
Flowering stages almost si.multaneously, 3 days ahead of unhardened plants in
s~.milar variants. Deepening of the tillering nodes was observed in
e~perimental plants even before the onset of drought (Table 2). The
+illering nodes are the principal organs in which new shoots form, and when
the above-ground organs and parts of the root system die, they can regenerate
entire plants. This property obviously has special significance to mass
death of plants in unfavorable conditions, particularly in droughts.
Considering that wheat plants with a deeper tillering node enjoy better
conditions for root system development and for maintenance of viability in
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` ~ ' c; `I;;~ "'t`' s
Effect of caryopses hardening and presowing treatment on
wheat grovrth in dry soil: Phytotron chamber, 1976. 1--
Unhardened (soil moisture content 70 percent), 2--hardened
(moisture content 70 percent), 3--unhardened (maisture
content 40 percent), 4--hardened (moisture cor,tent 40
percent), 5--unhardened, CCC (moisture content 70 percent),
6--hardened, CCC (moisture content 70 percent), 7-- un-
hardened, CCC (moisture content 40 percent), 8--hardened,
CCC (moisture content 40 percent)
Table 2. Depth of Tillering Node Following Drought Hardening of Seeds
Treated with Retardants and Grown in Soil With a 70 Percent
Moisttire Content; Greenhouse, 1975
Variant Treatment Tillering Node Treatment Tillering Node
- with ~p~h~ ~ Variant With Depth, cmn
Unhardened Hardened
1 Water 0 2 Water 15�O.G8
5 CCC 15�0.05 6 CCC 30�0.10
9. BES lOt0.06 10 BES 20�0.12
unfavorable environmental conditions, farmers usually try to plant the seeds
deeply (7-10 cm); however, deep sowing weakens the sprouts as a rule. To
the extent we know, causing the tillering node to develop deeper by presowing
light and high temperature treatment of sprouting seeds (19) has not enjoyed
practical application. We can see fram Table 2 that treatment of seeds with
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CCC deepened the tiller3.ng node of the plant by 15 mm, while BE5 treatmenti
deepened the node by 10 mm. Combination o� drouqh hardening of the plant
and retardant txeatment el.icited even greater deepening of the tillering
node--up to 30 mm with CCC and up to 20 mm with BES.
'rhe data presented here attest to the poaitive in�luence retardants have on
the depth of development of the tillering node in wheat, which was noted in
relation to CCC for the f irst time by Zadontsev et al. (16). A similar
ef�ect was observed for the first time when seeds were treAted with BES
retardant and the plants were drought hardened, and when drough~ harden3.ng
was combined with CCC and BES retardant treatment.
The tillering node of wheat in soil may develop deeper as a result of both
low temperature and moisture content on one hand and agrotechnical techniques
on the other, for example by changing the planting time or increasing the
seed planting depth. The latter, however,~may have an unfavorable effect ,
on caryopsis germination.
Deepeninc~ of the tillering node is an i.mportant ~actor of plant drought
resistance, especially in the early phase of plant development. When the
tillering node is located deeper in the soil, plants receive an additional
stimulus for growth; a larger tillering node forms, and the roots penetrate
more deeply in the soil, which promotes fuller utilization of the moisture '
reserves available in the soil.
Table 3. Change in Heat and Drought Resistance of Leaves in Response to
Seed Drought Harciening and Retardant Treatment (See Table 2 for
the conditions; greenhouse experiment, 1975)
Temperature of Cell Tuae of Leaf Cell Death ~
Variant Treatment Death, �C in Response to Dehydration, min
With
Tillering St~ Tillering Stem
Extension Extension
Unhardened
1 Water 58 74 95 100
5 CCC 60 77 105 110 .
9 BES 59 74 120 140
Hardened
2 Water 60 76 115 135
6 CCC 62 SO 120 150
10 BES 59 77 140 165 _
6
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Table 4. Effect of Seed Hardening and Retardan~ Treatment on the Structiuxe
of the Yield for Plants Grown in Dry Soi1 (Phyto~ron ~hamber, 1976)
BapHa?rr pcr�~iNll, KO1qC0~ KOJIOC OB ~ePeb Kol~iocaa, IOOO~epen,
, c?~ cM s?conoce r r
t8~ He3axa,7etittb~e
I. Bot~a - Ko~irponb (g 95 ' G,i i3,4 12,0 1,3 55,0
3, � ~acyxa ],0 73 4,2 ?,0 1,5 0,2 40,~~
5, CCC-tco~iTponb 11 i00 8,4 l2,5 l5,7 1,6 54,7
7, ~ 3acyxa 10 84 5,i 8,f 7,5 U,8 42,6
9, 6~C KONTPOJib 1Z 94 8,8 13,2 44,2 1,3 55,2
11, ~ aacyxa Lpy 82 5,4 9,8 8,0 U,9 45,i
(13) 3aKaneHHwe
2. BoAa KoFtrponb ~(9) 104 a,7 l3,8 13,5 1,3 53,2
4. ~ 3acyxa (].0) 89 4,8 7,8 10,0 0,8 42,8
6. CCC - KOtITPOJib ~,1) 98 7,0 12,7 20,0 1,5 57,0
8. � aacyxa ~ lp 85 5,8 10,0 17,2 i,i /i4,0
i0. 53C-KOHrponb ~,2 102 8,8 13,3 i9,? i,3 58,4
!2. s aacyxa Zp 88 5,9 l0,8 i7,6 1,2 52,1
Key:
1. Variant 8. Unhardened
2. Plant.. height, cm 9.' Water--control
3. Ear length, cm 10. Drought
4. Number of ears 11. CCC--Control
5. Number of grains per ear 12. BES--Control
6. Ear weight, gm 13. Hardened
7. Weight of 1,000 grains, gm
. r .
Table 5. Effect of Seed Hardening and Retardant Treatment on Relative Yield
From Plants Grown in Dry Soil (Phytotron Chamber, 1976)
Variant Grain Yield, Variant Grain Yield,
~ of Control ~ of Control
Unhardened Hardened
l. Water--control 100 2. Water--control 100
3. " --drought 15 4. " --drought 60
5. CCC--control 123 6. CCC--control 115
7. " --drought 60 8. " --drought 80
9. BES--control 100 10. BES--control 100
11. " --drought 70 12. " --drought 90
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Tab1e 6. Effect of Seed Hardening and Re~ardan~ Trea~cnen~ on the Structure
of thc~ Yield From Plants Grown in Dry Soil (Greenhouse, 1976)
- I~m+ ,,qneu oca, 4e onoa qi+c o aepeu I9cc 1 aepex' ,
n�p~~ cw cw Koe e Konocc e Konoce r
Ne38K8110HNb18 �
(8) 1. BoAa - Kourponb 84 6,4 !i 21 .98,3
(g 3. ~ - 3acyxa 48 5,4 41 1f 28,2
(],p 5. CCC xouTponb 83 6,4 43 24 40,0
(9 7, �--3acyxa 45 6,0 !2 ~3 38,1
(l~, 9. B~C Koiirponb 87 6,3 !2 21 /~0,7
(g 11, b--aacyxa 5i 6,0 12 !4 37,0
(1,2) 3aKaaeHe~te .
(8 2. Bo,~a-KOiirponb 85 8,6 !3 23 38,5
(9 4. x - 3acyxa ~i9 5,8 12 !3 40,0
(ZO 6. CCC - xoHrponb 84 6,3 i3 24 4i,2
(9' 8. aacyxa 48 6,3 43 16 41,0
(11 10. 5~C Koitrpo.7b 71 7,0 i3 24 38,4
{9)12. ~ --aacyxa 52 6,y 13 !8 40,5
Key;
l. Variant 7. Hardened
2. Stem length, cm 8. Water--control
3. Ear length,,cm 9. Drought
. 4. Nuitiber of spikelets per ear 10. CCC--control
5. iVumber of grains per ear 11. BE5--control
G. Weight of 1,000 grains, gm 12. Hardened
Determinations of plant drought and heat zesistance showed (Table 3) that
treatment of plants with CCC increases drought and heat resistance, while
treatment with the retardant BES raises the dehydration resistance of leaf
~ells more. As we hypothesized, the heat and drought resistance increasing
effect was more signi�icant with plants that were drought hardened and
retardant treated than with unhardened plants. Thus the results in
TaLle 3 indicate that complex treatment of the plant elicits an increase in
leaf cell resistance to high temperature and dehydration.
Soil drought initiated in the ear formation stage in the homeostatic
con3itions of the phytc~~r~?: had a strong influence on ear formation (Table
4). Ear length and t;~e number of snikelets and grains forming in the ear
decreased, having an effect on ear weight. The weight of.1,000 grains also
dFCreased. Empty ears and puny grains resulted in high yield losses. As a
rE:sult the loss of grain from unhardened plants (Table 5) was 85 percent.
7~e yield loss was lower when caryopses were subjected to retardant treat-
ment before planting. The loss due to soil dryness was only 40 percent
for unhardened plants treated wi.th CCC, and even lower--30 percent--for
plants treated with BES. Drought hardened plants experienced a 40 percent
yield loss due to drought. The yield loss was decreased even more by
combining plant hardening with retardant treatment. The yield loss of
hardened plants treated with CCC retardant was 20 percent, while that of plants
treated with BES Was only 10 percent (see Table 5).
8
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dus. ob~~rv~~ian~ r~howecl tiha~ ~o~.l drought led, among ~1an~~ ~hati w~r~ nc~ti
dr~ughe h~~c~~ne~,tio a gr~ati~r d~cr~as~ in sti~m growth ra~~, a d~er~ase in
~~r l~nqtih ~nd w~l.qhti, anc~ a drop in ~he number o~ spik~le~s per aar and tih~
nwnbar of gr~ins per ~pik~l~ti, in compar3.son witih harden~d plantis (sge
Tabl~ 6 ancl Fiqure). Hardened contirol and retardan~ tir~atied plan~e were
t~11~r ~nd ~he ear gra3n c~ntient was hiqher, as ~ resulti o~ whi.oh the plantig
_ suff~r~d l~~s frnm drough~ and produced better grain.
As wf:1 can se~ from th~~~ reaults, complex treatmenti of wheati before planting
r~is~~ th~ droughti and h~ati resistianca of tihe plantie. Apparentily ~he
in~~grati~d e~�@ct streng~h~n~ protec~3ve mechani~ma againsr tihe in~urioug
~etion of tiisgu~ dehydr~~ion.
BIBLIOG~tAPHY
1. G~nk~l', P. A., "Drough~ Resietance o� P1anti~ and the Wayg for Rai~ing
i~," in "Tr. In~tia fiziol. rastieniy AN SSSR" (Proceedings of tihe US3t2
Acad~my �Ynstitiuti~ of plan~ Physiology), Vnl 5, No l, izd-vo Nauka, 1946.
Genk~1', P. A., in "Fixiologiya sel'~kokhozyaystivennykh rasteniy"
(Physiology *~f Agricultural Plantis), Vol 3, 1967, p 87.
3. G~nk~l', P. A., in "Problemy za~ukhoustoychivosti rasteniy" (Problems
of P1anti Drought Resistance), Vol 5, izd-vo Nauka, 1978.
4� Genkel', P. A., FIZIOL. 12ASTENIY, Vol 25, 1978, p 889.
5. Badanova, K. A., and Tomakhin, N. A., S.-iCH. HIOL., Vol 11, 1976, p 31.
6. Tolbert, N. J. IIiOL. CH~M., Vol 235, 1960, p 4~5.
7. Leh, H.-0., ANGEW. BOT., ~~ol 37, 1964, p 212.
8. Catney, H. M., ANNUAL REV. PLANT PHYSIOL., Vol 15, 1964, p 271.
9. Halevy, A. H., and Wittwer, S. H., NATURh1ISSENSCHAFTEN, Vol 52, 1965,
p 310.
10. Halevy, A. H., Ashri, A., and Ben-Tal, Y., SCIENCE, Vol 164, 1969, p 1397.
11. Zadontsev, A. I., Pikush, G. R., and Grinchenko, A. L., "Khlor'1:holinkhlorid
v rasteniyevodstve" (Chlorocholine Chloride in Plant Growing), Izd-vo
Kolos, Moscow, 1973.
12. Kharanyan, N. N., FIZIOL. RAST:,'NIY, Vol 14, 1967, p 542.
13. Kharanyan, N. N., Burkina, Z. S., and C~seynova, G. M., FIZIOL. RASTENIY,
Vol 18, 1971, p 415.
9
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14. 8okar~v, K. S.~ Karanov, E. N., and 3vanova, R. P., "rnventor's
C~r~ifica~e No 365887, 28 Novemb~r 1972.
15. gokarev, K. 3~, Pxusakova, L. D., Murav'yev, S. A., and Chizhova~ s. i.,
F~~Z~L. RA3T~NrY, Vol 24, 1977, p 1078.
. 16. zac~ontsev, A. t., pikugh, G. R., and Grinchenko, A. L.,
"Khlorkholinkhlorid v rasti~niyevods~ve," Szd uo Kolos, 1973.
17. M~tskov, SOV. eOTAN., No 1., 1936, p 98. ~
16. G~nk~i', P. A., and Margolina, K. P., DOKL. AN SSSR, vol 86, 1952,
p 849.
lh. Kuperman, M., in "~izioloqiya sel'skokhozyaystivennykh rasteniy,"
Vdl 4, Moscow, 1969, p 7.
I
COPYRiGHT: izdatinl's~vo "Nauka", "Fiziologiya rastieniy", 1979
11004
~50: 1840
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~NZYMOLOGY UDC 57~.15.036
IN~LUENCE OF EFFECTORS ON TEIE FOLDING OF AN II~4ri08ILIZED PROTBIN (TRYPSIN)
Moecow MOLEKULARNAYA BIOLOGIYA in Rusaian Vol 13, No 3, 1979 pp 673-680
manuecript received 28 Jul 78
/Article by V. V. Mozhayev, K. Martinek and I. V. Berezin, Departmenti of
Chemical ~nzymology, Moecow SCate Univeraity imeni M. V. Lomonoeovf
[~ext] A study has been made o� the f olding of immobilized tryp-
sin in the presence of various effectore of its enzymatic
activiCy. The trypain~ coval.ently bound to Sephadex G-200
or Sepharose 4B, was folded in a concentrated solution of
- urea with simultaneoue eplitting of ite S-S bonda. with the
help of dithiotraeol. The preparation was then separated
from the eplitting agenta, one of the effectors of enzym-
atic activ~.ty of the trypain was added (boric acid, benz-
amidin, pancreatic and soy inhibitors of trypein, ethyl
ether of N benzoyl-L-arginine, methyl ether of N-tozyl-L-
arginine) and the reaction of the ~nnnobilized enzyme in the
absence of catalysts of thiol-diaulfide exchange was studied.
It was shown that the reactivation of the trypsin in the
presence of apecific aubstrates and proCein inhibitors
' takes place with the sams yield (2-SX) as in their abaence;
in the presence of benaamidin and boric acid, the reactiv-
ation yield of the immobilized trypsin is increased by fact- "
ors of 5 and 12, respectively, and amounts to 15 and 40X.
In a comparison of this :esult with the statistical prob-
ability of formation of 6 native S-S bonds from 12 SH-groups,
it is evident that even in the absence of a catalyst of the
thiol-diaulfide exchange (i.e., under conditiona of irrevers-
ible formation of S-S bridges) it is possible to increase the
reactivation yield of the enzyme several thousand times if
the enzyme ia immobilized on a carrier and is treated with a
soluble effector. A model is proposed according to which the
action of various effectors of trypsin on its folding is ex-
plained by their attachment to intermediately folded forme
of the protein and by a subaequent sh:ift of equilibrium
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~oR o~~zcinL us~ orr~Y
eoward eh~ "c~rrece" conformer~~ Accorfling eo ehi~
. a~~umption, ~v~n ~C ~ar1y ~e~~~s of folding, when 3-S
~ bonde have noti y~t bean formed, a drawing together eak~~
place of catal,ytically-ac~ive gYOUpe of serin~-hi~C3d~,ne
pair~ end n eorpt~.on sector ie organ3zed; f inally~ the
formation of ~n active canter of the ttyp~in (a draw~,ng
eogether of the caCalyric and sorption sectore) tiakes'
p1ac~ ae the la~e eCage of folding.
_ 'I'h~ problem eapir, b3ogynCh~~3~ of prote~.ns, a fund~mental taek of con-
temporary molecul~r b~.ology /1/. The queation has ae y~C noe been resolved
on how folding of g eynthesized polypeptidg chain into g funcCionally-
uctive protein takes place. To reeolve tihie it seemed fruitful to study
folding of prote~.ns in vitro /2-4/. It is no longer doubted thati the native
conformation of a protein hae been programmed in iCe initial etructure /3/.
How~ver, th~ m~chaniem~ which should function 3n the 11v~ng cell and pro-
moCe a mor~ effecCive (~.n particular, a faeter) fold3ng of proteine--ae
compared wiCh model experimenrs in vitro--ara not yet clear. It was ehown
earlier /5/ Chat a positiive role in the proceas in vitro can ne played by
natural immobilixaeian of the polypeptid~ chain on ribosomes. In the
present nrticle, a atudy has been made--on a model of trypsin, c~valently
conn~cted to Sephadex or Sepharoae--of th~ influence o~ various effectors
on ehe procesa of folding of an unfolded proCein (reactivation of the
~nzyme).
A multitude of inetabolitea are diesolved in the living cell. It ie well
known that some of them are capable of affecting the etructure and function
of proteins /6/, hence it ie completely probable that they can participate
in folding~ Actually, there have been isolated, from cells, compounds of
a 1ow-molecular (enzymes) /8,9/ nature, capable of effectively catalyzing
the thiol-disulfide exchange in proteins and substanCially accelerating
their reacCivation. NeverCheless, folding, in vitro, of eome enzymee into
the native conformation does not occur even under optimal concentrations
of catalysts of the thiol-disulfide exchange. Reactivation can be realized,
wit~ a large yield, onl~ in the presence of subetratee, cofacCors or euch
metallic ions (Ca2+, Mg Zn2+) without which enzyme functioning is not
possible. Thus, reactivation of takaamylase in the presence of Ca2+ /10,
11/, lactate-and malate-dehydrogenases /12/, glyceraldehyde-3-phosphate-
dehydrogenase /13/ in the presence of NAD, enolase, fumarase, aldolase aad
se-~~ral denydrogenases /14-16/ in the presence of their apecific aubstratee
nioceeds with yields excezding SOx. In the folding of heme-containing
proteins /17,18/, a necessary stage, according to Fischer, et al. /19/, is
c.,~rdination of the porphyrine ring of heme and (to a lesser degree) ao-
ordination of the iron atom.
These results /10-19/ indicate that, for aome proteins, reactivation with
high yield is possible only when the information about the native confotto-
ation, contained in its initial atructure, ia supplemented by the action
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of ~ffectore~ The mectian3~m of th~.s ~.nfluence upon Che folding of protie~n~
ean be represen~ed, accord~.ng to Teipel and Koahland /15/, ~.n Che following
way: in the fold~.ng of ~ polypepCi.de, ehera i~ �ormed both "correce" and,
a18o, "incorrect" intermediate conformer~, and tihese are ~.n g etiaee of dy-
ngmic equ~librium~ Additio~ of an ~f~fectior which reacts predominane~y witih
the "corr~cti" intermediat~ conformer ehould (~f the activat~.on barrie~ of
the e~ansieion beeween ehe ~oafor~~r~ i~ n~~ tiod gregti) lead to the desir-
~b1e shift of QqL~.1~,brium and, in the laeti analyais, ro an ~ncrease 3n
yield of the reacrivaeed protein~
The present article hae eeveral new ~~peces. ~irat, atiudy hag been made
of the inf luence of eoluble effectore on folding of ~mnobilized protein;
earlier /10-19/, for thie purpoae, only free en~ymes were used. The essen-
tial dieeinction in our work 3s that, earlier, in the enzyme which was un-
folded in solution by a d~naturing agent, Qith~r tihe S-S bonda were not
ep11t at aii /iz-19/, or, when they ware ap11C, react3vation was carried
out by addition of glutatione /10,11/. Our model (an enzyme ~mnob~.lized
on a carrier) makes it poesib~.e eo trace the reoxidation of the splir S-S
bonds even in the abgence of catglyata of the thiol-d3aulfide exchange, .
where the S-3 bridges are formed ~rrevereibly and, consequently, the inter-
mediately-forming "incorrec~" (catalytiically inactive) conformatione of
- the enzyme molecule are also irrevereibly fixed. As the reault of immobili-
zation, the yield of reactivated protein under these conditions*, which are
not favorable for folding, reaches eeveral percent /5/; this ie quite
enough to stay within the limite of the ueual methoda of ineasuring catalytic
activity. Second, atudy~has been made of the dependeoce of yield of re-
activation on the mechaniam (the method) of combining the effectoY with
the ~nzyme. Earlier /10-19/ it was ehown that an increase in yield of the
native form of protei~e during folding in vitro ia caused by the addition
of effectore ~hich aet~ basica~ly, in the finiahing atages of the procesa.
Thus, for example, in study of th~ intermediately-folded forms of taka~-
amylase, it ~aas found that the first 3 of the 4 native S-S bonda are formed
during the folding, independene of the presence in the system of Ca2+ ions.
HoWever, formation of the fourth disulfide bridge, and the etructural re-
- conatruction in the macromolecule, which lead to th~ catalyrically-active
conformation of the takaamylase, were seen only after addition of the Ca2+
ione /11/. On the other hand, it i~ known thate during renaturation of
carboxypeptiAase, the centers of cambining of the various effectors of
catalytic acCi.vity (in particular of inhibiCore) are formed at different
stages of the process of folding of the polypeptide chain /20/. Hence, it
ie of interest to atudy the influence of auch effectors, too, which might
react with the protein precisely in the early aCage of its folding.
~ In the presence of catalysts of the thiol-disulfide exchange (when the
"incorrect" S-S bonds are split, go as to form, in the end, the "correc~"
bridges corresponding to the thermodynamically stable, native conformation
of the protein) the yield of reactivated enzyme approaches 100X /5/.
13
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Influence of effeGeore of catalyti~.c activity of trypsin on reactivatJ.on of
tihe enzyme immob~lized on 3ephadex G-200.
20�, pH 8.0, 5 x 10'~ M enzyme, in tihe absence of catalyzere of the ehiol-
d~.sulfide exchange
~ 2 a~K,ocd .
3 neH~e~rn~ 5 m~r~euA M etiA 8 9
nopaMeT{+w peaKn~ua4xH 6e3 aQ,� aecKHA rH� coeO~rA ~,pNp N�6em ~~Q �ta 6eH~� 0op~r
� tAQNtbpO ra6xmp NHI'N01110p ~ONJI�L�9p� ~IIA�L�9p1'M� YNJLNN Itr~J10~~
TQHtlCIt118 ~NtlCN118 ~N~~ ~d~
KONCT3H18 ANCCOLlHBqNN ~ ,
Ii0F1D~1t'KC8 9f~1t~1CKTOPO C ~
l181NBN6tM cpep~~ee� 8~l0'~s 3~10'~ 8~l0'~ 10'~ !0`~
Toa~,11f - (29~ (30) 10'~ (SiJ� (32)� ~33) ~22~
KoHUenTpaut+a 3cpcplKTO�
pa, h1 i0's 90's i0'~�' 5�i0'~�� !0'= !0"~
- Adxo~t QeBKTNHBI(NN~ �,6 3 2-4 8�--5 2-4 2-5 !5 40
?
H 4pB11lNO! ~NBVlNYt! KOM4lMtP~WMHB V p 1 3O vc~e p~c~icoqa~ux~a~e)~i
Hri
,~rtpei~uuua 40-30~,~.
Key 1. Reactivation parameters.
Conatant of ehe diesociation of the complex of the
effector with the native enzyme, M.
Concentration of effector, M.
Reactivation yield, X.
2. Effectore.
3. wiChouC effectore; 4. pancreatic inhibitors of trypein;
5. soy inhibitors of trypein; 6. ethyl ether of N-benzoyl-L-
arginine; 7. methyl ether of N-tozyl-L-arginine; 8. benz-
amidin; 9. boric acid.
* Value of Michaelis constant, characterizing the form~tion of the
acylenzyme.
Initial value of the concentration; after 30 houra,
consumption of the eubatrate did not exceed 20-30X.
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u4D ' a , ~.4 ~r ~
~ ' ~ �
~ ~
~
~ ~ Yo a � ~ t ~ ,
~ ~ ,
' ~ ~
~ s
~ ~
~ ~ � '
r o
JO t0 JOvoc , ~0 t0 JOvice
Fig. 1 Fig. 2
Fig. 1. Growth of epecific activity cf trypein imQnobilized on Sephadex
G-200 during 3.te folding in the absence ~f catalyzere of thiol-diaulfide
exchange. 1.--without effector; 2-10' M benzamidin; 3--10'1 M boric
acid; pH 8.0, 20 5 x 10'~ M enzyme.
~ Fig. 2. Growth of epecific catalytic activity of trypsin, immobilized on
Sepharoae 4B, during its folding ia the abaence of catalyzers of thiol-
dieulfide exchange. 1.--wiChout effecCor; 2--10'1 M H3B03; conditione
ag in Fig. 1.
ordinata: enzyme activity, X; abeciesa: hours.
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In order to f~.nd euch ef�~ceore, we tie~tied a numb~r of compounde wh~.ch
C1~~rly d~.f�er in character of re~cti~on wirh the aceive ceneer of erypsin.
it wgs fouttd th~e boric acid, which is an inhibitor of alpha-chymotryp~in
/21/ ~nd of oth~r serine proee~.nases, includ~.ng tirypein /22/~ Eorme a com-
plex wiCh ewo carnlytic~lly-active gYOUps contained in the nucleoph~.l-
active center,--tihe OH-~roup of eh~ ~erine and the im~.dazo~. radical of
t~i~tidine /23/. Benzgmid~.n, a comp~titiive inhibitior of eryps~.n, reacts
~1~cCrogtigt~.cally witih Che a~paragine radical conCained in ehe hydrophobic
sorpe3on "pocketi" of tihe actiive center /24-26/. Speci�~.c substraees
(erhyl eeher o� N-benzoyl-L~arginine and me~hyl etiher of N-Cozy~.-L-arginine)
nre gtit~ched to th~.s same sorptiion "pocket" (electrostatically, with an
addieional hydrophob3c interaction), and also form, wirh the active center,
a hydrogen bond (with participation of the serine radical) ~24-27/; further,
in the acylenzyme there add~eionally appeara a covalent bond of ~he sub-
~erate radical wiCh the OH-group of the component nucleophil /24-26/.
Prorein ~nhibitore (pancreaeic and soy), which form complexes with trypain,
"close" not only the ac~ive ceneer but, also, the more importi~nt part of
tihe surface of ehe enzyme globule /24-26,28/.
PROCEDURE
We h~ve desCribed earlier /5/ the substiances and the meChoda of covnlent
attnctiment of erypsin to Sephadex G-200 and Sepharoae 4B used in this mork.
A typical experiment in atudy of the folding of a protein was the following:
rhe S-5 bonds in an immobilized enzyme were split, as described earlier /5/.
'Then the preparation was separated fram the splitting agenta and one of the
effectors of enzymatic activiCy of trypsin was added: boric acid (Soyuz-
reaktiv, USSR), benzamidin ("Sigma", USA), specific substrates or protein
tnhibitors of trypsin ("Reanal", Hungarian Peoplea REpublic), and reoxi-
~ation of the S-S bonds was carried out by the method described earlier by
~~s /5/. After a cerCain interval of time we Cook samples of the immobiliz- ~
ed trypsin and determined the enzymaCic activity (using the specific sub-
str.,Ce ethyl ether of N-benzoyl-L-arginine), following the familiar procedure
/5/.
RESULTS AND DISCUSSION OF THEM
Tt~e influence of effectors on reactivation of immobilized trypsin in the
absence of catalyzers of the thiol-disulfide exchange: the effecCors used
we-e added in excess with respect to the enzyme in concentrations signifi-
c~~tly exceeding (in the case of protein inhibiCors of trypsin and specific
i�nibitors) or commensurate (for low-molecular-weight inhibitora) with the
dissociation constants of their complexes with an active center of native
enzyme (Table). Like the protein inhibitors, the highly-specific substrates
have practically no influence on folding of the immobilized enzyme: the
kinetics of reactivation in all cases, is described by a curve of Type 1,
in Fig. 1; for comparison, the Table presents the corresponding magnitudes
of reactivation yield.
At the same time, low-molecular weight inhibitors significantly increase
the yield of reactivation of the immobilized enzyme--by a factor of 5 or 10
~6
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with addition of benzamid~.n (Fig. 1, curve 2) and of bor~.c ac~.d (curv~ 3),
reapecCively. A poaitive inf luence of these inhibitiore on reactivaC~.on of
trypain was found by ua wieh immobilizaCion of the enzyme not only on
Sephadex G-200 (Fig. 1) but, also, on Sepharose 4B (Fig. 2, data f.or H3B03).
From data on the depende.?ce of ~he yield of react~.vaCion of tiryp~~.n, ~mmor
b~.lized on Sephadex G-200, on concentration of the moat ac~ive e�fector--
boric acid (Fig. 3)--it appeara that lowering the concentration down to ~.0-6
M practically does not decrease the yield of reactivation of the trypsin
(30-40X). It can be aesumed that the conetant of combining the boric acid
with the fold~ng proCein ehould be of the eame order 10-6M). With
native enzymea--alpha-chymotrypein /21/, eubtilyain, alkalin~ peptidase,
_ pancreaC~c lipase, pencillinamidase /23/ and trypain /22/, the boric acid
attaches aubatanCial.ly worae (constanC of diseociation of the complex with
the active center ia 10'2 - 1 M). Explanation of the reasons of so great
differences in the combining of the Yigand wiCh native and folding proteins
(more than four decimal orders with reapect to the constant of dieaociation
- of the correaponding complexes) requires further atudy.
u
E 40 , o
e o o
~ o u
~ a~ ~ ��t
o y ��J
~ 20 ~ ~�r
~ ~ e 30 � �
a
~ ' - ~ i '
4 P. ~
-l9 ~H]g~.j /0 ZO JOvot
Fig. 3 Fig. 4
Fig. 3. Dependence of the end yield of reactivation of trypsin, im~obil--
ized on Sephadex G-200, on concentration of boric acid added during fold-
ing of the protein in the absence of catalyzers of thiol-disulfide exchange.
_ Conditions as in Fig. 1.
Fig. 4. Growth of specific catalytic activity of trypsin, immobilized on
Sephadex G-200, during its folding in the presence of atalyzers of thiol-
disulfide exchange (4 x 10'4 M of oxfdized and 4 x 10-~ M of reduced gluta-
thione) (1,2,4) and in their absence (3). 1,3,4--10'1 M HgB03; 2--without ~
effector. Conditions as in Fig. 1.
ordinates and abscissas as in Figs. 1 and 2.
17
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Re~ctiivation of inunobil~.zed trypsin in ehe presence of catialyzere of th~.ol-
disulfid~.ne exchange: ?f fo~.ding of irmnobil~.zed trypain is carried outi in
the presence of glutaChionea which cata~.yze thiol-disulfide exchange, the
yield ~E reactivation reaches almost 1009' even in the ~baence of effectiore
(due to one effect of the ~.nnnobilization) (Fig. 4,1) . As for the kinetiice
of the procese, reactivation in the presence of glutatihiones procaeda more
rapidly than in their absence (compare curves 1 and 3 in Fig. 4). Th~.s
agrees with the idea that the reaction rate ia limited by the chem~.cal
stages of the thiaL-d3sulfide exchange /7/.
Addirion of glutathiones at the end of the folding, carried out in the
presence of H3B03 (Fig. 4,3), ~eads to furtiher increase in catalytic act-
ivity (curve 4). This result 3ndicaCes that at~achment of the boric acid
to the folding proeein does not control the fixing of all of the native
S-S bonds; some of them are formed incorrectly. Since, in this experi-
q~emt, catalyzers of ~he Chio1-diaulfide exchange were added withouC addition
of urea (without aupplementary unfolding of the proCein) it can be thoughC
that the following redistribution of Che S-S bonds, accompanied by an in-
crease of caCalyCic activity (Fig. 4,4) does not require substantial con-
formar3onal changes in the protein.
Mechanism of the influence of effectors on the reacCivation yield in the
absence of catalyzers of the thiol-disulfide exchang~: The reaults ob-
t~ined can be explained in Che following way. In the early stages of the
folding, both "correct" and "incorrect" intermediary conformers are already
being formed. They are in dynamic equilibri~n, the mobility of which is
deCermined by the magnitude of the activation barrier. tnlith addition to
the folding protein of an eff ector able to form a complex only with "correct"
intermediary conformer, shift of equilibrium should take place to the aide
_ ~f increase in concentration of the "correct" conformer due to the free
ei,ergy of combining:
"incorrect" conformer ~ "correct" conformer ~
complex of the "correct" conformer with the effector
However, such a mobile equilibrium can be seen only at early stages of
fo?3Lng, whan the polypeptide chain is still suf�iciently mobile (in particu-
l~r, is not fixed by "inco�rrect" S-S bonds). In this case, i.e., in the
complex with an effector, the "correct" conformer, continuing to fold, is
fixed by the correct (native) disulfide bridges, as a result of which active
trypsin is formed with a higher yield than in a camplex without effector.
Surh a situation is realized, apparently, only in the presence of boric
ac'd or benzamidin. It can be assumed that these inhibitors positively
ir luence reactivation of trypsin, precisely for this reason, that they are
capable of combining with the folding protein even at the early stage of
the process. This means that even in early sCages of folding of trypsin
there are formed separate fragments of the active center which react with
the inhibiCors of cataly~ic activity, i.e., a drawing together occurs of
the catalytically-active groups of a serine-histidine pair with which the
boric acid reacts and a sarption sector is organized on which the attach-
ment of benzamidin takes place.
18
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Anoeher eitiuae~.on ~.s seen in addieion of epecif~.c eubatrgtes or protein ~.n-
hibitors. These compounda are ttble to react with euffinient ef�icacy only
wi~h a complere~.y-organiz~d active center. Moreover, formation of a com-
plex di the enzyme witih a proeein 3nhibitor, asaumes a definite organixation
of a rather large seceion of tihe surface of the globule near the active
cenCer. The fact tihaC these effectore do not influence the regctiivation
o� Crypsin ~Table) indicatea, evidently, thaC they are capab le of combining
wiCh inreXmediary conformers of the protein. This meana that formation of
rhe ~ct~v~ cenCer finiahea only at Che last ~tage of the folding. In tihie
case, those moleculea which have already f ormed the "incorrect" intermediaee
conFormationa (have been fixed by "incorrect" S-S bonds) cannot be recon-
atructed into correct conformers--tihis would require (in ehe absence of
caCalyzers of thiol-disulfide exchange) ~naurmounCably-large energy ex-
pendiCurea on apli~ting of covalent S-S bonds. Hence, effecCora, highly-
specific to Che aceive cenCer o� trypain, are not nble even to shift the ~
conformation equilibrium to the side of enzyme renaCuraeion.
Study of reactivaeion of immobilized enzymea in the presence of effectora
is important at the very leasC from three points of view. First, atudiea _
of the system model the folding o� proteina in vivo. 'I'he model suggeated
by us has mer3t because it takea into account that poly?peptides, in bio-
syntheais, are "immobilized" on ribosomea /1/. Trypsin, in this model, ~
is seen as "simply" a protein, ita specificity ie considered only in se1-
ection of effectors. It is knawn ChaC trypsin, in vivo, is not direc~ly
folded; it is formed by limited proteolyais of the inactive predecessor
Crypsinogen /34/. Neverthelesa, even for auch an "unfortunate" (from the
point of view of folding) protein, as trypein, it has been posaible in
thiy work to show that the immobilization and an effector, in their common
action, substantially increase the effectivenesa of reactivation. In com-
_ parison with the statistical probability of formaCion of six native S-S
bonds fram 12 SH-groupa (0.01%) the reactivation yield increased by a
factor of several thousand, reaching 30-40%. It must be emphasized that
folding of protein was studied by us precisely in the absence of catalyzera
of the thiol-disulfide exchange. Hence, it is possible that in living
naCure Che redistribution of S-S bonds in folding of protein does not
play so subsCanCial a role as could be thought on the basis of experimenta
carried out in vitro with ribonuclease /35/, lysozyme /36/ and Che pan-
creatic inhibitor of trypsin /37/. These proteins could be reactivated in
the free (not immobilized) form by a fundamentally different mechanism:
through intermediate products with a non-native bank of disulfide bonds
with their subsequent breakup in reactions of thiol-disulfide exchange and
formation as the result of conformation restructure of native S-S bridges.
As shown in the present article, it is possible by immobilization, with
mutual action of an effector, to completely avoid the intermediate form-
aicion of incorrect S-S bonds.
Second, it is known Chat some proteins, after unfolding, with splitting of
inCramolecular S-S bridges, are not able to renature /3/. It can be hoped
that, as the result of immobilization,it is still possible to bring about
renaturation of such proteins if the corresponding effector can be selected,
~g
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In solving thia problem, it is posaible to try ~o reactiv~te ~.rrevereibly
den~tured enzymes based on Che known achemme /38/. This would permit
mult~.n~e use of enzyme preparationa under technolog~.ca1 cond~.e~.on~ /39/.
mhird, sCudy of renatiuration of enzymes ehould reveal mechanisme of Cheir
denatur~Cion which have as yet not been diecovered. Thia has compllcated
creatiion of scientiific bases of seabil~.zing enzymes anci, consequently,
inhibies, to ~ definitie degree the introduction of biocatalysis into ,
prnctiise /39/.
~ BIBLIOGRAPHY
+ 1. Ingrem, V., Biosynthesis of macromolecules (in Rusetan). "Mir",
Moacow. 1975
2. Anfinsen, C. B., BIOCHEM J. Vol 128, 1975, pp 737-749
3. and Scheraga, H. A., ADV. PROTEIN CHEM. Vo1 29, 1975,
pp 205��301
4. Wettlaufer, D. B, and Ristow, S., ANN. REV. BIOCHEM. Vol 42, 1973,
pp 135-138
5. Mozhayev, V. V., Martinek, K. and Berezin, I. V., MOL. BIOL. (in
Russi~n) Vol 13, 1979, pp 73-80
6. Degli, S. and Nikol'son, D. Metabolic Pathways (in Russian). "Mir",
Moscow. 1975
~
Saxena, V. P. and Wettlaufer, D. B., BIOCHII~tISTRY, ~ol 9, 1970,
pp 5015-5022
8. Venetianer, P. and Straub, F. B., BIOCHEM. BIOPHYS, ACTA, Vol 67,
1963, pp 166-168
9. Givol, S., Goldberger, R. F. and Anfinsen, C. B., J. BIOY,. CHEM.
Vol 239, 1964, pp 3114-3116
].0. Takagi, T~and Isemura, H., J. BIOCHEM. Vol 57, 1965, pp 89-96
11. Friedman, T. and Epstein, C. J., J. BIOL. CHIIri. Vol 242, 1967,
pp 5131-5140
12. Chilson, 0. P., Kitto, G. B. and Kaplan, N. 0., PROC. NAT. ACAD. SCI.
USA Vol 53, 1965, pp 1006-1014
13. Deal, W. C., Jr., BIOCHEMISTRY, Vol S, 1969, pp 2795-2805
14. Teipel, J. W. and Koshland, D. E., Jr., IBID., Vol 10, 1971, pp 792-798
20
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15. � , , IBID., Vol 10, 1971, pp 798-805
16. ,~BID., vo~ ii, 1~972, pp 4100-4107
17. Harrieon, S. G. and Blout, E. R., J. BIOL. CHEM~, Vo1 240, 1965,
pp 299-303
18~ Babul, J. and Stel~.wagen, E., BIOCHEMISTRY, Vol 11, 19)2, pp 1195-1200
19. Fisher, W. R., Taniuchi, H. and Anfinsen, C~ B., J. BIOL. CHEM~,
Vol 248, 19~3, pp 3188-319S
20. Ko, B. P. N., Yazgan, A., Yeagle, P. L., Lettich, S. C. and Henkene,
R. W., BIOCHEMISTRY, Vo1 16, 1977, pp 1720-1765
21. Berezin, I. V., Kolomiytseva, G. Ya. and Martinek, K., DOKL. AN SSSR
(in Rusaian) Vo1 171, 1966, pp 1213-1216
22. Rotanova, T. V., BifuncCional reversible inhibitors of serine protein-
ases (in Rusaian). Candidate diasertation, Moacaw
23. , Vasil'yeva, N. V., Ginodman, L. M. and Antonov, V. K.,
BIOORGAN. KHTMIYA (in Russian) Vol 4, 1978, pp 694-698
24. Mosolov, V. V., ProCeolytic enzymes (in Ruasian) "Nauka", Moscow. 1971
25. Steitz, T. A., Henderaon, R. and Blow, D. M., J. MOL. BIOL., Vol 46,
1969, pp 337-358
26. Blow, D. M. and SCeitz, T. A., ANN. REV. BIOCHEM., Vol 39, 1910,
pp 63-105
27. Matta, M. S. and Stanley, A. D., J. $IOL. CHEM., Vol 249, 1974,
pp 732-743
28. Chotia, C. and Janin, J., NATURE, Vol 256, 1975, pp 705-708
29. Berezin, I. V., Kazanskaya, N. F. and Larionova, N. I., BIOKHIMIYA
(in Russian), Vol 35, 1970, pp 261-269
30. Harry, J. B. and Steiner, R. F., EUROP. J. BIOCHEM. Vol 16, 1970,
pp ??4-183
31. Inagami, T. and Sturtevant, J. M., J. BIOL. CHEM., Vol 235, 1960,
pp 1019-1026
32. Seydoux, F. and You, J., BIOCHEM. BIOPHYS. Res. Communs, Vol 44,
1971, pp 7451-7457
33. Mares-Guia, M. and Shaw, E., J. BIOL. CHEM., Vol 240, 1971, pp 1579-
1586
21
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34. Neurath, H. and Walah, K. A., PROC. NAT. ACAD. SCI., USA, Vol 73, '
1976~ pp 3825-3832
35. HanCgnn, R. R., Hanm?es, G. G. and Scheraga, H~ A., BIOCHEMISTRY,
Vv1 13, 1974, pp 342~.-3431
35. Acharya, A. S. and Tanuchi, H., J. BIOI~. CHEM. Vol 251, 1976,
' pp 6934-6946
37. Creigt;ton, T. E., J. MOL. BIOL. Vol 113, 1977, pp 275-341
38. Martinek, K., Mozhayev, V. V. and Berezin, I. V., DOKL. AN. SSSR,
(in Russian) Vo1 239, 1978, pp 483-485
39. � , in the book, Successes in bioorganic catalysis (edited by
Berezina, I. V. and Martinek, K.) ~in Rusaian) Izd-vo
Moscow State University, Moecow. 1978
Copyright: Izdatel'stvo "Nauka", "Molekularnaya biologiya", 1979
8586
CSO: 18~+0
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MARINF MAI~SAL9
vnc 59i,487:599.537
ON THF; PROBLEM OF CHENIORECII'TION ZN DOLPHINB
Kiev VF~BTNIK 200LOQII in Rues3~n No 3, 1979 received by editore 27 Dec 78
pp 3-11
,~Arti~le~ by a. B. Agarkov and 9. A. Q31.evich, Inetitute of 2oology of the
Ukrainian 58R Acadea~y of 3elenceJ
LTex~ The ability to differentiate ehemical. atimuli by means of taste and
smell 3g chara~teristic of ~ii m~~iB to a grester or lesser degxee. Rep-
resentetives of the Cetacee. order were coneidered Qn exception, be~ause the
peripheral sect3one and conduction paths of the olfactory ar~el,yzer, a8 ae11
se the gustatory pa~il.lae on the eurfsoe of the tongue, are redu~ed in ~hem
~Kukenthal u. a., 1 89; Ra~r~.tz, 1903; Sonntag, 1922). However~ observations
of the b~havior of dolphins under netural ~onditione and ocear~ariuma (Yab-
_ lokov, 1961; Lil~y, 1.965) made it neceseary to revise the Pormed opinion.
In connection aith the gbsence in Odontoceti of eome enatomical etructures
of the olfactory analy2er specialized exteroreceptivea perceiving chemic~l
stimuli ~an be only in their orel, cavity and, moat probably, in the mucous
membrane of the tongue. A. V. YQblokov (1957, 1961), describing the recesses
on the root of the tongue of the White Whale, characterized them as organs of
chemical sensation. Experiments conducted according to specie~~ ~ethods (Sokolov
et a1., 1971; Kuznetsov, 1974; 1978s; 1978~) he?ve ahown that dolphina can
differentiqte a number of subste~ncee diasolved in Water (indole, camphor, tri-
mett~ylemine, valerianic and caproic acide, secretion of ena]. and pro$tate
glands and so forth). It ahould be noted thet mar~y of the eubstancee very
aell perceived by dolphins are inadequate stimuli for the gustatory receptors
of the tongue of other mammals. Along With this, aubstances poesessing a
purPly gust~atory effect, srith the exception of bitter substances, aere poorly
distinguished by exp~rimenta]. animals. In connection with thie the opinion
(Kuznetsov, 197&S) tbat stimuli in loW concentretions are perceived by spe-
cial gustetory receptors belonging to the system of the trig~inal nerve xas
expressed.
,
~The report dQtQ were presented at the 3eventh All-Union Conference on Marine
Mauimals on 20 to 23 Septeffiber 1978 in 3imPeropol'.
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inv~~t3~~ting ~Ehe hieto~,ogy nf the mueous m~mbrane of the tongu~ of dolphina,
V. Ye~ 9oko~ov ~nd 0. V~ VoJ.kova ~5971~ eatabl3~hed tha~ the epitheZ~Q1 ~in-
3ng of th~ fleep caviti~e a~ ~he bound~ry between the root and body of th~
~on~u~ h~~ a di~tinctive etruoture ~ whieh makee it poe~ible 'EO B86L1n'l@ $hA'~
th~se formatione part3e3pn~e in the pereeption o� ehe~?3c~1 et3muli~ It w~e
po~~ibl~ to detect strueture~ eimilQr to gust~?tory paptl.~e~e end bulbd on the
bottom ~nfl in tihe 1,Qtera]. Walle of ~he abov~-mentioned esvities (9uQhawsk~~Q,
197~; Sucho~aek~~~ et al.~ 1q`~3; ~el'kov3~h eb a1., 1976; aiievi~h, i978;
l~hom~nko e~ Q].., ~.978)~ The ~bundanee and diversity of ~he intr~organ inner~
vatiion o~ the ton~ue of ~ooth~fl Whal,ea 3s ~n ind3reet proof of ita chemonen-
sory fwnction (~~ievsch, i975; vai3ui~n~? et Q1., i976). Thus, it w~s proven
th~b chemoreeeption oceupies an import~nt place in the vit~t act3vity of
toothe~ ahal~s.
Nev~rthel~ag, eome aepecta of the problem of chemoreception in cetaceane re-
quir~ a~urther etudy. The avQilabie morphological data Qre insufficient for
an expl~nation of the eharscteriatica of dolphine~ perception of varioue chem-
ical atimuli estiabliehed by exper3mentel investigatione. Furthermore, there
ar~ contradtctory opinSons concerning the cleesifi~ation oP ehemi~al senea-
tion in tioothed Whales. 3ome authors iden~iPy 3t tirith amell, ahile othera
conaider it e we11 developed een~e oP tas~e.
On th~ bueis of the ~bove-stated ae set for ourselves the task of studying
the micramorphology of tissue structures of the mucous membrane of the tongue
nf th~ bottie-nosed dolphin.
The investigation uas conducted on the tongues of 12 adults, 2 newborne end
3 9- to l~.-month old fetusea. The materi~l zras taken 1 or 2 houre after the
animals' death and Was �ixed in 12x neutrel formalln. Sicaining ~rith hemato-
xylin-eosin, carmine and alcian blue and impregnation With silver nitrate ac-
cording to BielschoWeki-Qros and Kampae Were used.
The mucous membrane covering the tongue of the bottle-nosed dolphin is amooth~
d~es not contain pQpillae so characterisLic of other mammals end ie closely
~oined With the underlying layers. A xeakly pronounced sub~ucous layer is
f~und only on the root at the transition to the orophes~ynx. The thiclsness of
the mucous membrane ig not the same in diPferent sectiona, increasing from
_ the top to the root, that is, on the top, 1.8 to 2 mm, in the middle of the
body, 2 to 2.3 mm and on the root, 2.5 to 3 mm~ Beaically, the mucous mem-
brune thickens at the expense of t. proprie. The epithelium covering the
lorsal surface of the tongue is Plat end stratiPied. The external keratinized
~.ayer is r~presented by three or four rosrs of P1Qttened cells ~rith dense nuc-
lel and the boundaries betaeen them are poorly discernible. The epithelial
lining oP the cavities on the root is formed by a amall number of rows of
cells with clearly differentigted, round nuclei. The keratinized lqyer ie
absent here.
Papillae in the tongue of the bottle-nosed dolphin are located at the bottom
Qnd in the lateral s~~alls of root cavities. These are protuberancea of the mu-
coue membrane, ahich do not rise above the surface of the tongue, because
24
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their height ~100 ~0 200 um~ iB smaller than ~he depth of the ~~viti~~~ In
~he ~ongue o#' th~ bottle-nosed dolphin ae dete~ted two ~ypes of papi~l~e--
fung3Porm ~nd con3cal~ The ~'ungiform papil~.Qe ~~ig. 1) are ~he largeet Qnd
- h~,ve a narroored ba~e and an expanded ~op. They exe of two types--with a ~on-
cave and convex ~op. The etroma of these papil~~e is ri~h in ceilular ele-
ments, Sm~ll secondary pap311ae penetre~te into the epitheliwn. 9ome fungi-
form pQpil~ae ere surrounded w3th a torue. The eonieal papil~ae ere em~ller.
Th~re ~e two v~rietieg of con3ce~.1. pap311~e--with Q pointed and rounded top
(fi~~ 2)~
` 4'', ,
1 !
~ Ny' ~
~ i 4 "�.~y . 1 . 1 .
f3t~~ t:' ~
F�4~A. y
~
t ~
L`A H
~
\~f
_ ,
. * 1~.
"
w~, i' ~*~i
,
i~ ~ N ~ t,~t ~t,y`,~~. i r,~ ~ ~
.
t : ~+~r y 1 ~ ' ~ � .l:
' _..�t,~ ' .',.t E fJ,~Lt� ~ 1` w. .
. .{q,' ' ~ _ t _ :'i ~ : '
. ~ ~ ~!~~Y~: y~ rS,,�
~ . ~i ^'`,~i.~f v~
~ ~ ~ + r; � � . . .
, ~ t';~
~A . 1
~ ` ~+~,w+ir'~' ~ .
�y~ ti �
`t'M`~'~ F�~ w t ,�'+1~~!~i
~
� ~
{r~. e . � 1* '~~,r.A~ f~ f: ~ ~ f,~ ` .
~ r ~ 1�~~*~'i4~'. .
~.,.~.~�f G _
g` .~r.~ri;
~i~.~.r1
Fig. 1. A Fungiform Papilla in the Tongue of the Bottle-Nosed Dolphin
a--hematoxylin-eosin (ob. 6.3, oc. 7); 6--impregnation according to Bielschow-
ski-Gros (ob. 20, oc. 10).
In addition to the described two types of pe,pillee in the tongue of the bottle-
nosed dolphin there are elongated ribbon-shaped papillae, as ~rell as papillae
of an incorrect form. Structures resembling taste bulbs are very rare. We
found them only in some fungiform pepillae (fig. 1~). They are bright forme~-
tions of a round or oval form. The gustatory canal is not quite clearly pro-
nounced in them e.nd sometimes it is not aeen at all.
Investigating the tongues of fetuses and newborns of bottle-nased dalphinsat our
disposal, we detected that their papillae are situated not only in cavities,
but also on the surface of the mucous membrane. They ere locelized (fig. 3)
in the area of the root and along the lateral edges of the boc~y, their number
ranging from 20-25 to 25-35� Some papillae can be grouped with fungiform pa-
pillae, while others, those surrounded with e groove and torus, can be clas-
sified as typical grooved papillae. Filiform pepillae are found occasionally
(fig. 4).
25
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, >ti~ ~
sx~
4 . Ft4~ X; r
~~~i.~. . i
~ ~
1 sy'.
~ S:
~~1 i
1
?
1 `
' ~ , ~
R
J
k~j
~'ig. 2. Conical Pepillae in the Root of the Tongue of the Bottle-Noeed
Dolphin (Hem~toxylin-~osin, ob. 10, oc. 7):
a--with a pninted top; 6--Witih a rounded top.
; ~ " J
Fig. 3. Diegram of Distribution of 'r s� r ~
Papillae Over the 3urface t
of the Tongue of Fetuses of
the Bottle-Nosed Dolphin: ' � s
1--large cavities e~t ~he boundgry :
between the body and root of the tongue;
2--papillae on the surface of the tongue;
3--root of the ton~we; 4--:rir,ge of tt.e 4
anterolateral edge of the tongue; 5--mu-
cous membrane of the lower surface of the
tongue. ~
In the tongue of the bottle-noaed dolphin there is a large number of glands.
They are absent on the top and along the lateral edges~ The bulk of the
glands is situated in the area of root cavities and behind them. Here they
form continuous fields divided by thin connective-tissue interlayers (fig. 5~�
The glands of the tongue of the bottle-nnsed dolphin are simple tubular-al-
veolar. The end sections of large glands consist of a large number of lob-
ules with narrow interlobular interlayers. 7'hey lie in the deep layers of
the mucous membrane. The main excretory ducts reach a considerable length
and their lumen is wide. Small glands consisting of tWO or three lobules
lie more on the surface. The excretory ducts of glends open into large and
26
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~m~11 n~?vi~iee on ~he Aurfaee of the tongue~ The g~sndular #'orma~iona in
the ~ongue ean be d3vided ~,n~o ~uo groups noti on1,y aecording to morpholog-
3e~1 ~h~r~etera, bu~ Q~.eo ~~~ord3ng to th~ nature of seeretion. The bulk
of tihe ~lande, when ~reated with aln3an blue, profluees a sharply pos3tive
re~etion to mue3n. The ema~,1, glande loeeted on the BurPaee ere not s~E~ine~
witih thia dye, Whieh m~ltes it po~eib],e ~o aBeum~ ~he pro~~in na~ure of ge-
cretion.
, ,
r
~
~�i ~ �'"'r
+st~ ~
t~ .
~ '
~,~~A ~t 1
.~~'_'` 0.03 Oe
curves 1-3 do not depend monotonously on maqnetic filed intensity: Some
of the points deviate from the trend. However, this deviation is significant
(98 percent) only at a magneti~: field intensity of H= 5 Oe.
- 38
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~l~ (2~ 3
J~octo~epuocr, ,qoero~epnoen j(ttC101lpNOCTb
~ bIWMNCIINa N NBdA OtKAOtIQNNq k~ pNBOA OtNAONI'NI1q NPIIS0~1
1 Ot ~r~ p Ot ~~00 ~1 ~ Of
0,0007 73,30 78,8T 8f,65
0,00f6 87,bb
~ 24?g4 25,88
0, 25 06,27 i2,7! 78,99
0,004 34,73 34,79 8g,2g
~
O;OtB 95,45 90;27 45;0T
O,OfS 100 89,i0 i00
0,05 198,9t !~'?3
0~1 88,07 99,97 ~~~95
.0;3 98,02 ~p~p,40 1~,95
0~5 '~,73 83~85 93~i2
99~95 i00
0;~5 ~97~55 i00~95 !00
8E3~84 i00 97,63
2~5 80,8! 88,27 93,27
~ 98,88 i~,40 99~94
4 99,98
5 99~24 99~24 99,97
8 1~'27 1~,88 l~,23
~~5 97,75 f00
Key:
1. X, Oe 3. Si.gnificance of deviatiion of
2. Siqnificance of deviation of curve 2 froan 1.00
curve 1 from 1.00 4. Siqnificance of daviation of
curve 3 fran 2.00
Concurrent research on the effect of magnetic field orientation (the seeds
were oriented with the embryo pointing north, south, east, and wast in
each Petri dish) on wheat caryopses showed that a positive effect does occur
in all seasons except spring (both stems and roots qrow more vigorously
when the embryo is oriented southward), being 8-10 percent on the average.
In sprinq the magnetic field orientation effect chang~s sign to equal
(-3) -(-4) percent. It should be noted, however, that there are great
difficulties in studying the dependence of the magnetiic field orienta~ion
effect on ~gnetic field inte:~sity, since the measurements must be highly
precise; it should also be noted that the fiqures presented above are not
precise.
BIBLIOGRAPIiY
1. Krylov, A. V., and Tarakanova, G. A., FIZIOL. RASTENY, Vol 7, 1960,
p 131.
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2. Tarakanova, G. A., "Efgeat of Constianti Magnetic Fi~lda on Plan~ arowth
and Enargy Ma~abolism," Cand~,8a~e Disear~ation Abstiracti, Moscow, ZFR,
1971.
- 3. marakanova, G. A., in "Ma~~rialy =Z Vges. aimp. 'Vliyaniy~ yeg~~~tv~nnykh
i glabykh isku~ativennykh magni~nykh poley na biologicheskiye ob"yekty
~ (Proce~dinga of tihe S~cond Ail-Union Symposium "~�fec~ of Na~ural and
We~k Ar~i�3cial Magna~ic Fields on Siological Ob~ects"), Selgorod, 1973,
p g8.
4. Navitskiy, Yu. z., Strekova, V. Yu., Tarakanova, G. A., and Prudnikova,
V. P., in "Vaprosy g~natiologii, r~diobiologii i bioloqicheskogo
d~ys~vl.ya magnitinykh poley" (Problema in Hema~ology, Radiobiology,
and th~ 8ioloqical Actiion of Magnetiic Fielda), zxd-vo Tomsk un-ta,
1965, p 329.
5. Str~kova, V. Yu., Serdyuk, L. S., and I~ b~dev, V. A., 3n "Materialy iY
Vses. s1mp. 'Vliyaniye yes~es~vennykh i slabykh iskusstvennykh
maqnitnykh poley na biologicheakiye ob"yekty'," $elgorod, 1973, p 111.
6. Novitskiy, Yu. i., in "Tez. dokl. sove.shchaniya po izucheniyu vliyaniya
magnitinykh poley na bioloqicheskiye ob"yekty" (Abstracts of Reports at
a Conference on Research on the Effecti of Magnetiic Fielda on Hiological
Objectis), izd-vo Nauka, 1966, p 50.
7. Tarakanova, G. A., Strekova, V. Yu., Chertok, N. 0., Novitskiy, Yu. I.,
and Bokarev, N�, S., "Materialy ii Vses. soveshch. po izucheniyu vliyaniya
poley na biologicheskiye ob"yekty" (Proceedinqs of the Second All-Unfon
Cnnference on Research on the Influence of Magnetic Fields on
F3ioloqical Objects), Moscow, Izd-vo VINITI, 1969, p 217.
9. Dul'binskaya, D. A., ~I2IOL. FtASTENY, Vol 20, 1973, p 183.
9. Novitskiy, Yu. I., and Travkin, M. P., "Materialy nauchn.-metod. konf.
'Khimiya, botanika, zoologiya (Proceedinqs of the Scientific
Conference "Chemistry, Botany, Zoology"), Belgorod, 1970, p 73.
10. Pittman, V. J., CANAb. J. PLANT. SCI., Vol 42, 1962, p 430.
11. Draganets, Ye. A., and lravkin, M. P., "Materialy II Vses. simp.
'Vliyaniye yestestvennykh i slabykh iskusstvennykh magnitnykh poley na
biologicheskiye ob"yekty'," Belgorod, 1973, p 161.
12. Nikulin, A. V., in "Materialy III Vses. simp. 'Vliyaniye magnitnykh
poley na biologicheskiye ob"yekty (Proceedings of the Third All-Union
Symposium "Effect of Magnetic Fields on Biologfcal Objects"),
Kaliningrad, 1975, p 18.
40
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13. Nov~.ti~kiy, Yu. t. , and Markma?n, Yg.~ V. ~n ~'Me~erialy zz Vses. simp.
'Vliyaniye y~~~~~tvennykh i elebykh iekute~vennykh maqnitinykh polsy na
b~,ologicheekiys ob"yetky'," Heigorod~ 1973, p 92.
14. Nnvit,~kiy~ Yu. t., anc~ ~s~~rova, o. Y~., in "Ma~erialy zz Vses. soveshch.
po izuoheniye vlayaniya magni~nykh po].ey na bioloqicheakiye ob"yektiy,"
Moscow, zzd-vo VtNZTI~ 1969, p 162.
1S. ~rik, E., and Torp, M., "Pribory diya nauchnykh 3eaZedovaniy"
(Znstrumen~s for Saientifia Research), izd-vo Mir, Voi. 40, 1971,
p 21.
16. Bogatina, N. Z., Verkin~ B. i., Kordyum V. A., Kordyum, Ye. L., and
Litivin, V. M., UOKL. AN U33R. 3ER. B., No 4, 1978, p 353.
~ COPYFUGHTs ~zdatiel'stvo "Nauka", "Fixiologiya rasteniy", 1979 �
11009
CSOs 1840
~
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SCIENTI9T9 AND SOIENTIF'r0 ORGANIZATZONS
SECOND ALL-UNZON' CONF~RENQE ON PROBL}~ IN BI4ME(3HANICS
R~.ga 1?~KHANIKA KOMPOZITNYKf{ MATERIALOV in Ruealen No 3~ 1979~ pp 560-56,2
[~tiele by I. Y~ Knets]
`Text] On the initiative of the Saientific Ooun~il of the USSR AN ~ca-
denq? of 3ciences] on problems in biemeohanlca, the Latvian 3SR Acaden~y of
3clenQes and the ?~atvian 53R Mlniet,ty of Health~ the Seaond All-Union
Conference on Problems in Biomeehanica was held in Riga from 18 to 20 April
1979~
Three hundred sixty seven xorks of 3oviet and 19 worka of foreign saientiste
wer~ selected by the orga,ni~ational committee for presentation at the
conference~ A11 the works xere grouped by 54 separate themea for each of
which a survey report was presented~ In makirag the selectSon the organi-
zational committee gave preference to xorks in which fundamental or prac-
tically important problems of applied biomechanics xere examined.
At the first plena.ry session four aurvey reports xere heard. Y. K. Kalnber~'
report examined current problema Sn medtcal biomechanics associated with
the ~pplication of compression-distraction apparatus in treating bone defor-
ma;:ions, in osteosynthesis and in restoration of ~oint funetion. In V. I.
Shum~cov's report the current status of development of the artifYdal heart
wa~ analyzed in detall. Different desSgns for its power supply and regula-
tion xere examined, and the biomechanical properties of the blood circula-
tion system xere evaluated in order to improve the construction of
artificial hearts G. A. Lyubimov's report xas devoted to a nex and rapiclly
~ieveloping area of biomechanics--r~sgiratory mechanics. The main problems
a;soc3ated xith development of an effective mathemat,ical model of respiration
~.'are investigated, and the influence of the mechanical properties of the
~~spira+,ory organs on the biomechanics of the respiratory process xea
evaluated. In his report~ A. P. Gromov gave an account of the basic aspects
of biomechanics in head traumas and evaluated the infl uence of not only the
mechanical but also ~Dhe,mox~phogeometric propert3es of the ekull on the char-
acter of injuries to it.
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~OR UF~aIAL U~~ ONt?Y
S1x ~ur+?~y ~portg w~~ ~ivan on fundam~nt~. probl~m~ of biom~ch~n~.~~ of
compaat bon~ tis~u~. In hi~ 3roport I~ V~ Kne},s ~xamin~d d~formation ~.nd
' d~~t~uctien o~ ~om~et bone tiseu~ from th~ vi~wpoint of th~ maohaniae of
~ h~rd~ ,d~fo~ii,~~ boc~y. Th~ d~gree of ~,te ansso~opy and pt~yaiaal non-
11ne~~ity w~ra ~v~uat~d~ ~nd qu~~tion~ of the viseoel~~~Ei~ b~havior of
bon~ ~isgu~ w~r~ pres~n~edt the ro1~ of th~ pie~oelectric ~ff~ct in bono
Na~ ~naly~~d~ E~ P. Podru~hnyrak'~ r~po~ was devo~~d to g~n~ra1 question~
of ~h~ agina of bon~ ti~~u~ ~nd th~ morpholc~gioal and biom~ohanioal ct~an~e~
in bon~ ~~oci~t~d xith tt~i~ proc~~a ~ In hi~ rsport~ G~ Q~ T'.faf~od ~v~.u-
at~d th~ ~ff~et of ~ diff~rent typ~ of influena~--~hiah inc].ude~ redueed'blood
~mpply~ N~i~htl~~~nes~ anrl ~adiation--on the mecha~nical prc,~~rti~s of `
human Hnd anim~,7. bon~ tissue. In 9~ S~ Tkach~nko's report current p~oblem~
in th~ f~~tion of ~g~n~~raterl bone in fu~3on of f~ac~ureg wa~ ~x~ml~i~d~
~h~ in#'~u~nae of ~lee~ric pol~,ri~atson and local m~ahanical vibration on
tt~i,~ proc~s~ xa~ ~v~7.uat~d, and the d~gres of change in th~ acoustio and
m~chani~~1 properti~~ of bon~ du~ing ~gen~ra~ion wag analy~ed~ A. K.
Muy~huli~' r~port N~ d~vot~d to aum~lex qu~~tiong of biom~chanic~ i~n
t~ating f~atur~by int~rnal ogt~osyntheai~~ evaluation of ~110- and
xendgeno~ bon~ supportg and ~tee1 pins~ In I. K. Yi1ka'g report biomechani-
aa1 probtem~ ari~ing flwcing the period nf rehabilitation w~rg examined~
in d~tail. Th~y ~ related ~o teaching o~ proper-wal.king~ compensatto n
for th~ motor flefeet and optimie ation of c~ontrbl of~ movemarrt~.
Six survey reporta Nere presented on questiona in the biomechanlcs of
~ndoprosth~tica. In Kh. A. Yanson'~ r~port general biomechsnical aspects
associated With endoprosthetics xere examined and the f~a,sibillty of
applying crystal insulators as atructural biomaterial~ for ondoprosthes~s
xas analy~ed. In Ya. M. Shersher's report the durabillty and strength
of endoprostheses of different 3oints Here evaluated. Ya. B. Kutsenko's
report xas devoted to evaluation of thQ positioning~ form and orientation
of different bone elementa in the ~joint by means of roentgenograpt~y.
In his repo~t V. S. 3hargorocLskiy anal,yzed the basic biomechanical principles
for treatment of articular pathology of the loxer 11mbs~ Such current
problems as determination of the biomechanical criteria for restoratim of
the functions of the knee ~oint depending on the method of plastics of the
ligamentous apparatu~ and the coxofemoral 3oint follo~~ing ~rthroplasty srere
examined. A. Revenko's report xas devoted to examina~,3.on of di:ferent
statoc~ynamic indices 1n orthopedic diseases and traumas of the loxer limb.
B. S. Rozenshteyn gave an account of the general biom~chanical criteria
for atanding and t+alking f'o~k~lox~,ng endoproathet~c~ of the knee
or coxofemoral ~oint and gave an evaluation of the electric activity
the ~high follo:.ing endoprosthetlcs of the knee 3oint using Sivash's method.
Three survey r~ports xere read an the biomechanics of the vertebral colwan.
V. Ye. Raykhinshteyn presented a reviQx of data on deformation and destruction
of the glemants of the spin~l cwlwnn--the .ver'~ebr~o., the intercertebra~.
discs and the dura mater spinalis. Analysis of the statics and c~ynamics of
the intervertebral discs under the inf`luence of different loads was presented
and the ~�eight-bea.ring capacity of the human spdnal column and vertebrae.xas
eva;luated. In Ya. L. Tsiv'yan's report the results of biomechanical
~+3
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~va~.ua~tion o� i;he meohantigenesis of ~ha ver~tebre~. oo~umn ware p~ea~nted
and th~ problams of ~txea~ing its poa~-~aumatia and degenera~tive deformation~
w~~~ di~~u~sed~ Biom~chanioal ori~eria were propoAed ~or aelecting the
optimal v~s~a.nt o~ bon~ plastios of ~Ehs ver~ebral aolwnn~ V~ E~ Belen'kiy'a -
r~porb w~ devoted to problema of the biomeohanios of ~taech~ndgeneeis
and ~roblems in treating gco~.iosis~ ~nolud~.ng the biomechanical basis for
op~rations on ~he interver~ebral d~.scs and bhe ver~ebrae in aco~~.oais~
New models of ~biologic~l. sys~ms` and me~thoda of investigation. in
biomedhanic~ w~r~ exami.ned in ~wo eurvey report,s. A~ S~ Vi~en~oi~ reported
on the principles of the physical model.ing of the elementa of pa~tho~
logical ~ait~ presented`a model of the elactriaal resistance of bone
~.isaue, evaluat~d in~uries' ~o the vartebral colu~nn ,from the pez~pective
of biomechanica ~nd examined ~hs functiona of ths foo~~ In R~ A~ Gurevich's
report th~ qu~ations of the investi~ation of the biome~hanical properties of
~;he lower limbs of man by ~the methods of vibration 3experiments and u].txa-
sound~ measurement of the presaure between bone.fragment,~~ increas~.ng
the information content of ~tatographic invegtigations and application of
qual~, t$t3ve electromicrograpt~y were exami.ned.
One survey report touchod on ~ene~l questions of biomechanics and zoology.
The oper~ker~ S~ F. Manziy, analyzed in detail the properties of deformation
nnd destruction of bone tissue~ ligaments and musclea of animals~ took
not,ice of the charact~r~.st,ica of the statolocomotion of the locomotive
apparatus of terr~strial vertebrates and evaluated the patterns of change
in the e~,ructure of bone tissue of animals during ontogenesis~
Ten reports were presented on the mechanics of soft biological tissues and
their substitutes. In his interesting survey of the mechanics of skeletal
muscle L. V. Nikitin examined different mathematical models of muscle, _
analyzed the deformation of muscle as a function of its activation and
evaluated its v3scoel~sticity. The sub3ect of artificial muscle~ was
discussed separately. V. I~ Yorob'~v's report explored the biomechanical
problems associated with the structure and mechanical properj~irs of or~e of
the b~asic components of biotissues--collage.n; the influence of its atructural
organization on the mechanical behavior of collagenous fiber was ~~"aluated,
among other things. Yu. Zh. Saulgozis' report was devoted to the ~naracter-
istics of deformation of skin~ the sclera and other types of soft tiss~les and to
analysis of the influence of different mechanical conditions on the regen-
eration of soft biotissues. R. P. Kikut examined the biomechanics of the
1~.rteries and tissue of the bra.in, paying part~icular attention to thrombo-
~enesis in the cerebral arteries. In his report V. V. Dzenis not only
ina~yrzed tn detail the characteristics of the transmission of ultrasound
~o the sotmd biotissues but also presented interesting data. on the applica-
tion of ult.ra,sound for characterization of coxarthrosis and fractures of
the bones of the knee and thigh. V. A. Kas'yanov gave a concise account of
the characteristics of the mechanical behavior of the ma~or blood vessels~
He reported on the most general mathematical models of these vessels which
describe deformation of them under different types of inechanical influences
and presented ex tensive experimental material. In V. Ya. Isanov's report
44
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an analy~is of ~he g~n~ra1 problems of ~he mechan~,oa of the n~roo~rdium was
g~.v~nt among o~he~ thinge mathema~ioal model~.ng of con~trac~tion of the
n~yooardium~ ~h~ v~.s~oelag~tia behavi~r of aaxd,~.ac muso~e~ and the meoha,nical
proper~iea o~ ~he ven~trioles of the heaa~^~ were desoribed~ N~ 8~ Dobrovaya'a
interea~ting ~port was devoted to prob].~ms of practioal ~,mportanae ~.n
3,nve~~tigat~,on of the propertiee of the meoha,i~i.aa1 behav~.or of ~the valves o~'
~the heart and the orea~ion of pros~thea~s of themt an integral evalua~ion
of ~the l~yc~,~codyn~mi.c ~.nd9.aes o~ the valvea was given~ and the dynami.as of
their op~ra~.on were analyzed~ B~ A~ Purinya's report was devo~ed to
questiona of selec~ion of su'bsti~utes ~or ~t,he b].ocd vesaels ~ ahe examined
the feasibi~i~ty,of wsing xenotransplanta and synthetic prostheaea for
replaaemen~ of in,~ured veasels ~ In A. M. Mo.vshov~.ch and N~ S. Gaveyushenko's
report the role of the 3nfluence of aggressive bioenvironments on the mechan-
ica1. properties of different polymeric implan~t,s was evaluated~ The au~thors
preaented a ma,thema~tical mode~. of the dea~rue~ion of implan~s in the bio-
environment and gave an evaluation of the stareng~h and antifriction proper-
tiea of bio3.nert thermoplasts for endopros~hetics of 3oin~ss~
FYve reports were presented on the meohanical propertiea of biological
fluids and the problems of biomass exchange. V. A, Levtov analyzed in
detail the cha,racteristics of the aggregation and orientation of erythro-
cytes~ compared the viscos3ty of blood in healtt~y and sick people, evaluated
the influence of additives on the indices of the dynamics of circulation
and examined the sinovial flow through the capillaries at different rates
of displacetaent. Y. M. Zaiko's report wa,s devoted to questions of the
hydroc~ynamics of circulation. A mathematical model of the movement of
blood in cavities with irregular geomet,ry and in a tube with distorted
wa11s was exa,mined in detail, and experimental data on blood flow in the
venous bed was preaented. Problems in the biomechanics of microcirculation
were interpreted in S. A. Regirer's report; he gave a deta,iled review of
the current status of the problem ~,nd analyzed a number of problems~ such as
mathematical modeling of ~}rogenetically active blood vessels, investiga-
tion of the isometric contraction of smooth mwscle, modeling of vaecular
tension. M~ A. Khanin's report was devoted to questions of modeling
the vascular bed and evaluation of the peripheral resista,nce of the
vessels. V~ N. Akimov analyzed problems associated ~th extracorporal
metabolism of biological substanc~s, chiefly with artificial blood circulatiian.
Twelve reports were presented on questions in the biomecha,nics of movement
' of humans and animals. In his report V~ Y. Beletskiy gave an account of
the principles of modeling bipedal. ~ralking and examined control of the
inertial leg during the phase of transfer and parametric optimization of
bipedal walking. Ye. A. Devyanin's report wa,s devoted to problems in
modeling of hexapocbus walking. In it the timing of the forces in .the legs
of the walking apparatus was also evaluated, use of a range-finder in
order to control ia,7.king robots during movement on complex sites was ~
analyzed and a model of an integral walking appara,tus was pr~sented. In
M. Y. Kudzyavtsev's report the results of experimental irvestigation on
the locomotion o~ humans and animals were reported, an evalua,tion of the
c~ymamics of moviements using special counter-comput~r systems was given,
. 45.
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and the diatr~.l~u~ion of ~he key reaotions in a system o~ the exoekele~on
type wa~ oxam9.~led ~ A. Ye . KobrSnsk~.y' a report was devot~d to th~
b~.nmechanics of workors' movemante--the ques~ion~ of th~ ma~ximum c~nduxabin
loads nn the lwdy of a pet~on ue~.ng a pro~eative aasembler's b~1.t~. '
~h~ mobility of human h~.nds and ~,mprovSng preoision movements in you~hs
with ~raining. V. M~ Za~.v~.orskty examined the problems of biomechanical
an~,lys~.s of human locomotion a~d gave an evaluation of the racLi.oisotope
method for determining biomechanical oharacteristics of aegmen~ts of the
human body. In his report V~ K~ Bal'tsevich gave an evaluation of the
external and interna]. determination of the paxameters of the development
of locomotive systems during human ontogenesis and a description of the
integral cr~.teria for evaluating the level of aahievement o~ the ob~ec~ive
in sport movements~ A~ N~ Laputin's report was devoted to problems of
developing technica]. means for shaping movements using automated contrc,l
in tea~hing ~thletes'motor training. S:'M~ Kozlov examined the
characteristics of muscle acti.vSty in differen~ athletic movements.
I. P, Ratov gave a concise s~atement of the principles of contro111ng'
the movements of athletes~ discussed spectral analysis of movement and
modeling of the processes of control of movement by ~the biomechaxiical
components of man. V~ T. Hazarov's report was devoted to questions of
the synthesis of athletic movements and vibrostimulation of muscles in
training ath].etes. F~ K. Agashin gave an account of problems of wave
biomechanics and gave an evaluation of training on biomechanical machines~
In the report of the Polish sc3entist E~ Maxynyak gene'ra1 problems of
ma.thematical modeling in biomechanics of mov+ements were exarained; among
other things, the da.ta of estimation of the dyna.nd.c properties of a.
person jumping with a closed parachute were,presented.
Five reports were devoted to questions of the biomecha.nics of pros~hetics.
Yu~ V. Kurochkin ga,ve an account of the biomechanical a~pects of st~nding
and walking normally and with different pathological deviations, presented
biomechanical criteria for the effectiveness of operative treatment of
children with congenita.l. dislocation of the femur;and ~,nalyzed the char-
acteristics of thE statics and kinematics of patients with compression
fractures of the vertebral column. V. I. Filatov's report touched on
q~:astions of kinematics not only o.f the upper limbs but also of prosthetics
for them. The author presented the theoretical grounds for developing
multifunctional prostheses for the hand and ~ve an evaluation of the use
of kineplastic tunnels in control systems for them~ In I. A. Mendelevich's
re~~ort the biomechanical aspects of creation of designs for prostheses and
m~asuring equipment were discussed, including ways of normallzing the
lunctional chaxacteristics of the knee joint of thigh prostheses and questions
-f the testi.ng interlinking surfaces as a way of rein forcing the joint. .
V. A. Berdnikov's report was devoted to questions in prosthetics after
amputa.tion of the lower limbs and a.nal~CSis of biomechanical designs for
constructing prostheses. I. Sh. Moreynis examined questi~ns of human
sta.tics and kinematics after prosthetics, gave an evalua.tion of the
stability of 'human orthograde posture normally and on a prosthesis and
investigated the influence of the normallzation of the position of the
OTsM [,expznsion +~nkno~m] on the results of p~asthet3:cs.
46
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A~t the end of ~Ehe oonferenae's work~ a meet,ing of e11 five seotions of ~he
9oien~.i.fia dounoil of the USBR AN on problems of biomeohani.oa was held~ in
wtiioh the re~ults of the conferenoe were disauesed and proposals ware made
for improving the oonferenoe's work in ~he futut~~
In the conolud,ing plenary eeesion Dootor of MedScal Saienae V~ 8~ Gurfink~l' ~
the chairman of the aeotion on ~he aontxol. a~d regula~tion of biol.ogioal
~ys~emes Doa~or of Teohn3oal Saienoe I~ V~ Kneta~ the chairmax? of the aec~ion
on ~he biomQOhanias of biological mater~als and syatemss Asaooiate Member
of ~he USSR AMN CAcadentiy of Medioal Soienaea] Y~ K~ Ka].nber~~ the chairman
of the section on medioal biomeohanica~ Doctor of Medioal Science V~ I~
Filatov, the ohairman of the seo~ion on the biomeahanics of substitut~s
for biological tiasues~ organs and syatems~ and Doator of Pedia~rios V~ T~
Nazarov~ the ohairmaxi of the eeotion on aport biomeohanlas gave a ehort
evaluat~on of the works preaented on the themea of their sections~ A
reaolution was made conoerning wider development of reaeareh in the main
trends of biomechanics
COPYRIGHTi Izdatel'etvo "2lnatne", "Mekha.nika:kompo~itr~ykh materialov"~
1979
93+30
G'~ 0 r 1840
47
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SOIENTISTS AND SQIFN~F~IO ORGANIZATIONS
RESOLUTION OF THE SEOOND ALL-tiI~lION C(~!iFBftENCE ON PROBLEM9 IN BIOME%INArRC3
Riga MEKNANIKA KOMPOZZTNYKH MATERIAI,OV in Russian No 3~ 1979~ P 563
["Text" of reaolution by the Seaond A11-Union Conference on Problems
in Biomechanics held in Riga on 18-20 Apri1 1979J
[Text] Three hundred ninety eight ~pecialiats took part in the work of
the second A11-Union Conferen~e on Problenus in Biomechanics. Among them
were 113 doctors and 176 n~stera of science. The number of those wishing
to participate in the work:of the conference was conaiderab~ly greater~
S urvey reports represented by the countryr's moat prominent apecialiats
reflected the basic accom~lishmenta of the different branches of biomechanics
not ~nly 3n the world in ge,~era]. but alao in our country in particular~
Durittg the per3.od between the First and Second All-Union Conferences on
Problems in Biomechanica (October 19?5-Apri1 1979) $iKnificant $uccess
wa.s achieved in the stuc~y of the mechanical behavior of biological tissuea,
systems and fluids~ in investigation of the processes of' regulation of
biomechanical systeaos~ in the biomechanics of artificial tiasues, organs
and systems~ medical biomeahani.ca~danalyais of control of movementa in
labor and sports. The number of dissertat~.ons and publications on the
bas:.c problems of biomechanics and the practlcal signific2~nce of
proposed scientific developments has increased.
In terms of organization the most aignificant event xas the creation of the
Scien tific Council of the U33R Acaden~y of Sciences on Problems in Biomechanics
xhich carried out work on detezmining the basic trendss in the development
of biomechanics and coordination of theoretical and experimental reaearch.
At the present time research in biomechanics is being conducted in ~ore than
30 institutes of the US3R Acade~qy of Sciences, the USSR Acadetqy of Medical
Sc'.ences and the union republic academies~ including the Ukrainian~ Latvian,
Georgian and Armenian SSR academies~ in 50 universities of the country~ 40
scientific research institutes of the USSR a~nd ~he uni'on-republic miniatries
of heal th and others.
At the same time the res olution of the First A11-Union Conference on Biome-
chanics on including a"biomechanics" specialty in the List of Specialties
48
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8oientifYc Workera remaina u~,~~,iiea~ ~ There ia �no~ one aingle pu~Sis~hed
organ on biomeohanios, ~nd speoialis~s axe stil]. no~ being txained widely
enough in the higher eduoational ~.ns~ti~u~ti.ons o~ ~the oountxyr ~ ~
Talcing the above ~.nto aocoun~t~ ~the oonferenoe reodmmends i
1~ ~ha~t~reeearoh in the basia ~trenda of biomeohani.cs be ati11 further
expanded~ paying primary ~t~ten~tion ~o ~t,he complex soluti.on of the'
mos~ urgen~ theore~i.ca1 problems and practioal taska=
2. ~that th~ Sta,~e Qommittee of ~the USSR Counoil of Mtn~.aters on Scienae
and Technology be requeated to int,roduce a"biomeahani.os" speoia].ty to
the 11et of speoial~ti.ea of acient,~.fic warkers ~ a step wh3oh would
fac3litate preparation of highly quallfied soientific personnel and at
the same time would insure higher quallty of ~the scie~tifda developments
accompllahed by themt
3~ ta,king into acoount the aoute need for a publiahed organ on biomechanica~
that the bureau of the Sc3entific Oouncil of the USSR AN on Problems in
Biomeahanice be requested to examine the question of the fea,sibility of
iesuing a apeciall~ed collection on biomechanics and to negotiate with
with the responsible o~gana for creation of iti
4. that the miniateries of of higher and secondaxy speciallzed education
and the USSR and union republlc ministeries of puhlic health be requeated
to support the initia~i.ve of the VUZ of the country in training
specialiats in the ba.sic trends of biomechanics.
COPYRIGHT: Izdatel'stvo "Zinatne"~ "Mekha.nika kompozitr~ykh mateMalov"~
1979
9380
CSO: 1840
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PUBLICATIONS -
CHEMICAL PREVENTION OF RADIATION CONTAMINATION ~
Moacow KHIMICHESKAYA PROFILAKTIKA RADIATSIONNYKH PORAZHENIY in Russian '
1979 signed to press 24 Jan 79 pp 2,3,4, 190.
[AnnoCaCion inCroduction and table of contenta from book by A. S. Mozzhukhin
and F. Yu. Rachinskiy, Atomizdat, 2550 copies, 190 pages]
[Text] This book, the first edition of which was issued
in 1964, is devoCed to medical prevention of acute
radiation sickness caused by external x-ray, gamma- and
neutron radiation in fatal quant3tiea. The questions of
research, experimentation, and the mechanisms of the
effect of radioprotectors are examined. Particular
atCention is devoted to sulphur-bearing compounds, but
data are presented on all classes of adequately effec-
tive anCi-radiation chemical compounds.
The book is intended for radiobiologists and pharmacolo-
gists, and for speciaLista in medical radiology, radia-
tion hygiene and protective medicine.
Eleven drawings, 56 tables, 422 titles in bibliogr~phy. -
In*.roducCion
At the present time chemical prevention of radiation contamination (chemi-
cal protection) is a recognized branch of radiobiology and radiation medi-
cine and no longer needs special ~ustification as it did 10 or 15 years
ago [14].
~he terms "chemical prevention" and "chemical protection" should not be
~aken literally. Pharmacological substances that, when introduced before
irradiation, completely avert the characteristic changes in the organism
known as radiation sickness are still unknown. The terms "chemical pre-
vention" and "chemical protection" should be taken to represent a concep�~~
tion of 3ncreased resistance or of decreased sensitivity in animal or human
organisms to the harmful effects of ionizing radiation. Future research
will demonstrate which of these two concepts most accurately reflects the
facts. For the time being an equal sign must be placed between them.
50
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}{iataric~iiy, th~ l~.r~~ work eo ~how Ch~ po~eibii#.ey ~f ehamica~.ly pr~-
venting r~diat~rni dam~ge in mammal~ aae that o~ Pate ~385~ ~nd Crdnkie~
~295, 298J, who d~mon~ernted in 1949-1951 ehat cyetein~ and glutarhione,
wh~n intr~du~ed ineo th~ organi,ems of miea and r~ee b~for~ ~~tha~. x-r~y
irradiation, ware cap~b],~ of pravene~.ng death in ~~ignific~nC number o�
animala~ However, the protective effect wae not conetant, whi,ch pro-
vnked ekepriciEm about the very Eact of protection. The final answer to
the queet3on about the possibi~ity of chemical prevenCion was left to
- Bacq (279~, who showed in 1951 that cyaCeam3na and cygt~m~ne, intYOdue~d
tnto mice before irr~d~.ation, permiCted 100~ eurv~val of the experiment~l
, animals, wh31e the controL animals euffered 100X mortality~
Redioprotective compoun8s were found in varioue claesea of chemic~l sub-
~tances, but the most likely onee for future uee were aminothiole~ end
their derivaC~ve~ (18,41J. But Che relaCively h~gh toxiciCy and narrow
therapeut~c range of the aminothioles and moat of their derivatives
cnu~ed dnubt as to the poss~bility of the~r usef ulnesa in protecting
humans from the harmful ef�ect~ of. ionizing radiation (230J. In the
succeeding years rhousands of new compounda were synrhesixed and etudied
in tesCs on mice, and gmong these effective r~dioprotecrive agenea were
discovered with a wid~r therapeutic range than that of cyaCeamine and
cygeamine; these finds stimulated intereat in the queetion (181~204,263j.
T'he moet thoroughly studied radioprotectanta have begun to be uaed in
x-ray therapy and oneological practice.
Atter the release of Che first edition a greaC deal of material was puU-
lished on new radioprotective agents and the mechanisms of their pharma-
cological and protective actic~n [18, 70, 103, 173, 183, 184, 256J. This
spared ~he authors of thie book the need to syetematically set forth the
prablem of chemical prevention of radiation contamination. In the pres-
ent edition, as previously, the superior radioprotective effect of amino-
thioles and their derivativea is examined, as well as their conversion in
the human and animal organism and their influence on the condition of the
organism. The suthors have had much personal experience in the synthesis
and analysis of these compounds. They also considered it essential to
compare the radioprotecCive effect of aminothioles with that of other
- radioprotective compounds, and to present new data on the effectiveness
of radioprotectants under different conditions of the action not only of.
x- and r-ray irradiation, but also of neutro+ns and protona, and data on
the effectivenesa of radioprotectants in combination. At the same time _
the authors have shortened or deleted several chapters and aections whose
contents fell outside the framework of analysis, testing and evaluation
of rFdioprotecCants.
51
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Table o� Cont~nts
IntrnduCtidn 3
i~ Chemic~~. properties o� radioproCectiants b
1. prop~rt~~g of r~d3oprorect3ve ~ubstance~ b
2. Mech~ni~mg uf ch~m3ca1 prot~cC~.on of organic ~ubatances 6
3. 3~Leceion of radioproCectante ~.n tegt mod~ls 24
li. Chemical protecCion of mice from the effecte of x-ray and
~-irradiation ~n the m~.nimum abeolute lethal dosage 48
1. protectinn oE mic~ by Yadioprotectante o� dif~erent
rl~s~eg nf chem3cal compounds 49
2~ Protect~on of mice by r~dioprotectanCa in different
combin~Cion~ ~ 75
~It. Chemic~l prdCecCion of variou~ mgmmalian species 78
1. prorecCive effects o� radioproCectanta in varying condi-
tiong of introduceion 78
2. ~ffectg of radioprotecCants with sublethal, fractionated
~nd long-term general irradiation, and aleo with local
x-r~y and a^-irradiation a~
3. Proteceive effect of radinprotectant with neutron and
proton irradiaCion 93
IV. Behavior of ~ulfur-bearing radioprotectanta in animal and
t~uman organiama 101
1. Absorption of radioprotectants 103
2. Distribution of radioprotectants in the organism 106
~ 3. Conversion of radioprotectants in the animal and
human organism 118
4. Itemoval of radiorrotectants and products of their
breakdown from the organism 124
V. Influence of r~dioprotectants on the functional state of the
organisms of hig~er animals and humans 128
1. Influence of radioprotectants on the metabolic process 129
2. Changes in the main physiological functions of the
organism under rhe influence of radioprotectants 134
3. Influence of radioprotectants on the aystems of nervous and
and humoral regulation of physiological functions 148
VI. Conclusion. Possible mechanisms of the protective effect of
chemical agents for prophylaxis of radiation sickness 154
Literature
COPYRIGHT: Atomizdat, 1979
9354 52
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ro~ o~~~cz~. us~ oNLY
PUBLICATIONS
CNEMICAL ANU BIOLOGICAL AOENTS ~OIt PLANT ~t~OT~CTIONt A~HO~T l~AN~~OOK
Mo~cow Kt~IIMICHESKIYE I BIOLOGICHESKIYE SREDSTVA 2ASHCHX7.'Y RASTENIY (KraCkiy
Spravochnik) in Ruseian 1978 signed to preea 7 Jul 78 pp 2, 4-6, 207
~Annot~C~.on, introducCion and table of contenCs from book by N. V. 3azonova,
"Ko1oe" Publishers, 100,000 copiee~ 207 pages]
[Text] Thia book conta~n~ ehorr iteme about preparationg
recommended for practical uae and experim~ntal producCion~
ttecommendations are given for the uae of cl~emical an8
biological proeective agents for agr3cultural cropa against
pestg and disease, and the conditione and baeic regulations
for their application are indicaCed. A lieC of prep,~rations
for protecCing plants in collective and individual gardens,
and the rules for their use, is presenCed. Safety measures
for working with peaticides are set forth.
The book is intended for plant protection apecialists,
agronomists, and aookhoz and kolkhoz directors.
InCroduction
In view of the further intensification of agriculture, protecting plants
for pests, disease and weeds is an integral part of a progressive technology
of crop cultivation.
The most important tasks involved in perfecting and r~inforcing plant
protection in this country were determined in the basic directives for
national economic development in the USSR for 1976-1980, accepted by the
25th GPSU Congress: furrher expansion of pesticide production, with
deliveries to agriculture of up Co 628,000 tona (in conditional ~~�r~its) wae
called for, as was the need to protect tfie bioaphere from p~llc;cion. Tl~e
July (1978) Plenum of the CC CPSU also devoted coneiderable attention to
the protecCion of plants~
'Che importance of the chemical method of protecting harvesCs from plant
pests and disease lies in its higfi effectiveneas, return for money expended,
and accessibility for use. For example, in 1975, Chanks to the use of
~ pesCicides to combat pests, disease and weeds in the main agricultural crops,
~ an additional 16.5 million tons of grain, 11.7 million tons of vegeCables
and potatoes, 16 million tons of sugar beets, 2.2 million tons of raw cotton,
3.6 million tons of fruit, 2.3 million tons grapes and other products, were
harvested for a total of 7.8 billion rubles.
53
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'Ch~ nh~micai meehod i~ now being perfeceed ar~th tihe a~m of r~duci,ng eh~
Jang~r of ~h~~~ prepar~eion~ eo hum~na, ~seful organi.em~~ ~nd narur~ a~ a
whal~. 'To ehi~ and p~re~.~eent pYepar~ttone are ~eing ~eplaced by lese
rp~i~rant Chemical~, qu~.ck to break down withouti tioxic re~idue ge wp11 e~
by 1~~~ toxic ~ub~tanc~s. Pr~paratione conraining arsenic~ mercury and
fluorin~ d~ri.vleivee are being almost totally eliminared t~rom Che aseorem~nt
nt Chemi~al~; n~w ch~mic~1 subseances harml~s~ to humans and usEful animal~
~r~ b~ing ineroduc~d, and Che forms and metf~ode of th~3r preparation are
b~ing p~rfec~~d.
In r~cent y~~r~ def~.nitie ~ucc~s~ ha~ been observefl in decreasing pe~ticid~
toxiri~y Co warm-blood~d organiem~. A large number of new peaticides
have be~n geudied ~.n d~eail, permitting �undamental revieion and renewel
of eh~ avai].able assortment of pesCi~:idee, thanks to the etforta of the
A11-Union ~cienCific res~arch 3.n~titute for hygiene and Coxicology of
p~g~icides, polym~r~ and plaetics, the All~-Union ecientific reserach
in~ei~ut~ for chemic~l mean~ of plant protaction, an8 o~her ~cienCi.fic and
production ~eeabli~hmenrs of the Minietry of Agriculture, the Minf~tYy of
Chemic~l Indu~tiry and Ch~ Mfniaery of Health. In 1965 the average toxicity
nf in~ectoacaricidee for warm-blooded organiema was 200 mg/kg, and itt 1975,
g~0 mg/kg, so the toxicity of the preparationa has been decreased almoet
Eiv~-fo1d. U~ring ehis period all preparatione that are highly toxic for
wgrm-blooded animalg were eliminated, as were resistant cumulative sub-
~tnnce~ that can cause chronic poieoning. As replacemenCa for DDT and other
chlorine organic compounda, phoephororganic compounds of inedium or low
roxicity for warm-blooded animale, which break down in one vegetation period
co nontoxic meta~olitea, were adopted.
'Che State Commission for Chemical Substances for Combating Pests, Plant
Uisease and Weeds recommended for agricultural use organic phosphorue
compounds ((antio], basudin, [bromophos], [gardonaj, ~volathonJ, DDVP,
� (amipho~sJ, metathion, (phosalon~, [B1-58j, (phthalophos] and others),
c~rbamic acid esters (sevin, [pirimor]), organic halogen compounds ([dilor],
[tedton], keltan, gamma-isomer HCC11 and orhera), nitro derivatives (acrex)
an~ others. About a third of the recommended preparations are selective
inse:ticides and acaricidea with a lethal dosage (causing over SO percent
mortality in test animals) of over 1,000 mg/kg. Specifically, this figure
is 1,900-5,000 mg/kg for ~gardona], 5,000-9,000 mg/kg for [dilor],
5,000-10,000 mg/kg for (tedion] and so forCh. The uae of these preparations
permits control of the chemical method and decreased danger to useful
mert.bers of the agroceuoeis. For example, specific acaricides and aphicides
(acrex, [tedion], kelt~n, (plictran, pirimor] and others) are 500-1,000 times
mo~e toxic to pests than to their main enemiea (chrysopids, coccinellidae,
pt~latory bugs, flies, etc.). Insecticides such as [gardona, dilar, amiphos]
are 100-500 times more dangerous for harmful insects than for entomophages.
New forms of preparations and methods of their application have been worked
out to Purther increase the efficiency and safety of chemical substances.
Preparations in granular form are highly effective for combating pests
(wireworm, grain ground beetle, weevil, cutworm, nematodes); they are also
significantly less dangerous Co useful organisms and contribute much less to
54
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th~ c~nCaminar~.nn o~ the envi~on~ec~t~ For wide~pread uap granu~ated
pr~pnr~eion~ b~~~d on Che gamma-i~omer HCCH, basud~n (va~.ekeon~,
" m~Chald~hyde, (phoephamid~, chlorophoe~ ~rc~, ere recommended~ Aer3a1
u1Crn-low volume ~praytng is being ~eeted for use in combaCing liarm�ul
luci?~~H, ,aeink bugs, end grain cutworm ne~ng epecial forme of Chese
~~r~rarat3on,
'1'he assdrtmettt o~ fungicides and seed mordante has been i~prov~d ~nd upd~e~d.
On green plnnr~, in~Cead of nordeaux liqu~d, 1ow-tioxia preparations ralated
to rh~ thiocarbamates and phthalimide~ (zine6, ~polycarbaxine], Cpolymarz~.na],
(polyhdmj, CdiCan M-45j, c~proxan, captan, [phthalan]) are us~d again~t
false mildew fungi and leaf epoti. To ~ucce~~fully combat p~,~dery mildew,
which affeces various agricultural cropa, ~caratan)~ acrex, and (moreseanJ
ar~ reconunended in addition tu colloidal sulphur. In recent yeare sysCemic
fungicides have come widely into use that are ef~ective aimultaneously
ggainar ~everal diseas~~ ahich may develop on the sgme cu].ture (benomil,
[uxg~n], [eopgin-M], etic.). It hag also 6een shown that such eyetemic
fungi.cide~ a~ benomil, [eopsi.n-M] and [uzgen~ may aleo be used a mordant~
for ~~~de and goil preparations.
Gre~e achievements have also been made in the aearch for non-mercury
bearing mordanta. Among the effective replacementa for granosan, mercur-
t~exane, and mercurbenezene are benomil, [vitarax], hexathuiran, ~tigam],
[EP-2] and others.
- Various biological preparations based on bacCeria and fungi are a part of
Clie broad practice of crop protection.
'Che use of selectively toxic rodenCicidea has noticeably increased. In
Eield conditions the Soviet preparation gliftor, which is practically
harml~ss eo birds, is widely used, and synthetic blood anticoagulants are
highly effective in combating rats and mice.
The entire contemporary arsenal of chemical substances for proCecCtng
plants has been thoroughly sCudied at the correaponding scientific estab-
lisliments of the USSR riinistry of Public Health, and based on this study
standards have been determined for the the use of these preparations:
optimal norms for expenditure of pesticides; permissible maximwn residue in
food products; permissible amount of the preparations in the air of work
zones; waiting period from last processing to fiarveat.
Successful solutions to the problems of regulating the chemical method
and eliminating the possible negative consequences to the use of pesticides
(such as poisoning of humans and domestic animals, of bees, which pollinate
plants, and of enromophages; formation of resistant populations of harmful
insects, acauds, and pathogen; desposiC of preparation residues in harvested
products, and pollution of the environment with these residues) may be
reached not only by selecting new preparations, but by rational and expert
inclusion of chemical methods in the zone system of protecting agricultural
55
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~OR O~FIC~AL US~ ONVY
Crop~, eh~ �ound~tion of whicR ~hould be eg~otacbn~cal prophylactiic
m~ellednio~y, A~horough analy~i~ of g11 blolog3cal pecul3ariete~ o~ th~
~~rnt~c~tad c:rep'd agroc~noeie ie pes~ntial~ and muet tak~ in~o nccount eh~
eC~nomie thr~shold of hgrmfulnese for eveYy pe~t epeeieg wieh3n the zonal
~~eCor.
13ranch ~nd zon~ insGituCee are Btudying and refining Che optimum at~ndards,
eime pariod~ and application methode foti the recomm~nde8 prep~rat~ons.
Knowl~dge and observ~nce of eha regulation~ e~~a6liehed for use of chemical
gub~t~nce~ 6y workers ~ngaged in plane protection and by agrochemical
kolkl~ox and ~ookhoz pereonnel sC~ould permiti a11 positiive aspecte o� active
~1~emic~l protecCion of plants eo be maximized, and to a11ow the elimination
df m~ny undeairable conaequenGes.
The pr~g~nt t~andbook is a practical guide to rhe exieting regularions for
ehe nppli.cation of contemporary pest3cides and their effective and eafe uae.
('I'ext] Table of Contenta
IN'TEtOnUCTION Q
CN~MICAL ANU BIOLOGICAL 5UBSTANCES FOR PLANT PROTECTION 7
Inaecticides and acaricidea 7
Mordants 19
Fungicides 24
Nematicides 29
Rodenticides 30
BiopreparaCions 32
A1dAN5 OF PROTECTING PLANTS FROM PESTS AND DISEASE 34
Grain crops 34
Legume crops 51
5ugar beet 58
Fi6er crops 68
I'erennial grasses 78
PotaCoes 85
Vegetable and melon crops 90
~ Oil crops 111
F'ruit crops 114
Berry crops 142
Grapevines, cit~us crops and tea 156
Tabacco, coarse tobacco, hops 169
INSECTICDES TO COMBA~T PESTS IN GRAIN AND GRAIN PRODUCT RESERVES 176
U5~ OF RObENTICIDES 179
56
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7UBSTANCE3 ~OR PROT~CTING PLANT9 FROM PEST3 AND DISEA9E3~ PEItMETTED ].90
FbR Pt1~LIC 3ALE AND U3E ~N COLL~C~IVE AND INDIVInUAL OARDENS
COMPAT~~ILI'I'Y OF INS~CTICIDE3, ACARICIDE3 AND FtJNGZCIDE3 198
ZONES OF EXPERIMEN'TAL PRODUC~ION U3E OF CHEMICAL StJB3TANCES FOR 202
pit0'T~CTING PLANTS
SAFETY MEASURE3 FOR WORKING WITH PES'TICIDE3 204
COPYRIGH~t Izdaeel~etvo "Koloa," 1978
9354
CSO: 1840
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PUgLICATI0N5
GLINICAL METHODS, DIAGNOSZS AND TREATMENT FOR ATTACKS BY TOXIC CKEMICAL AGENTS
Mogcow KLINIKA DIAGNOSTIKA I LECHENIYE PORAZHENIY OTRAXLYAYUSHCHIMI VESHCNEST-
VAMI in ~uesian 1978 signed to presa 26 May 78 ~~p 2-5, ].76
(Annot~tion, IntroducCion and Table of ConCente from book by Yu. N. Seroykov,
Meditsina Publishers, 110,000 copiea, 176 pages~
CTextJ 'fhis book is devoCed Co questions of clinical procedure, diagnosis
nnd ereaCment for patienrs affecCed by chemical agents in the event of mass
contamination of given areae by chemical weaponry. A ahort deacription of
chemical weaponry and ita featurea is given and the range of inedical assist-
ttnce, both 3n Che center of chemical contamination and at the atage of inedi-.
c~l evacuation, is set forCh, as applied to Che medical service aystem of
Civil llefense. The principles of antidote therapy which is of great impor-
tance in the treaCment of contamination by toxic chemical agenta~ are
presented. MeChods of self- and mutual assiatance, which to a aignificant
degree deCermine the course and outcome of contamination, are examined, and
the basic methods uaed to restore reapiratory function are elucidated, as are
~,~eans of individual and collective antichemical defense.
mhe book is intended for middle medical workers; it contains 17 drawings,
and 23 titles are included in the biography.
The high contaminating effect of toxic chemicals ~J wa$ irrefutably
demonstrated during the FirsC World War when the use of chemical weaponry
caused the deaths of huge numbers of people and serious damage to Army
personnel sub~ected to such attacks. In the following years it was found
that people who had been seriously affected by OV frequently became invalids
requiring constant medical observation or long-term treatment.
At the Genoa Conferettce in 1922 the question of forbidding chemical warfare
was brought up on the initiative of the Soviet Union, but the repreaentativea
of the imperialist states declined to discuss the issue. In Geneva in 1925
' an agreement was worked out forbidding the use in war of "...asphixiating,
poisonous or stmilar gases and bacter~iological substances." Although the
Soviet Union was among the firat to ratify this agreement, a number of
countries, including the USA and Japan, refused ratification. In 1952 on
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rhn initiueiv~ n~ ehe Sov~.gC Union rti~ Un3Ced Natione called for the rari�ica-
einn nE rh~ G~nevn protocnl by gll CountrieB~ Only in 1910 d~.d the U~S~
gdv~rnm~ne propog~ parei~l recognitinn of Che Geneva protocol, k~eping for
it~~l~ Che righC eo use nV not mentioned in the document~
Aftpr World Wgr II g coneinu~l ~~grCh went on in the U.S. and Che NATO
eouneri~~ for n~w OV, disCinguighed by highly contgmingeory features. Some
of Chese were ue~d by Che U~5. army in rhe war againsC Che Vietnamese p~ople.
Th~ grmie~ o� 3mperialise etaees are equipped wiCh exCremely eoxic OV, which ,
Cnn b~ caC~gorizpd in the fol.lowing basic groups: fatgl OV (~'PS, yperiees);
rhoge cgueing eemporgry incapgciey (paychomimetic, some irrieants and tear-
gas~s); and those causing short-Cerm incapacity. Foreign miliCary specinlists
ConCittually gtress eh~t the greaC effecCiveneas of OV ia manifested par-
ticulnrly clearly in ~udden attacks and when Che populaCion is inndequately
rrainpd in antichemical defense. There ie no doube whutever Ch&C the effec-
Civeness of chemical w~aponry is sharply decreased when prevenCivc and trent-
menC meagure~ nre taken cnrrectly and in time against OV affection.
The ttireat of mags conCamination of the population in the event that gn
aggressor employ~ chemical weaponry forces the Civil Defense (CD) medical
services to face a number of tasks, of which the most important are:
n) implemenCing all-round preventive measures with Che aim of avoiding OV
contaminaCiott; b) taking correct and timely measures to provide self- and
mutual assistance, and CD assistance in the chemically conCaminated zone;
c) to warn of Che laCer development of after-effects characteriatic of
seriou~ damage by OV. Although these tasks are faced in the event.of a
ChreaC, not only of chemical, bur of nuclear and bacteriological warfare,
the scope and nature of the medical measures taken in a zone of chemical
cnntamination are unique and specific. We have devoted much aCtention in
Chis publication to explaining Che peculiarities of damage caused by chemi-
cal weaponry.
The fulfillment of the Casks set before the CD medical services requires of
the middle medical workers a clear understanding of the nature of the medi-
cal measures taken under the threat of a chemical attack, in the zone of
chemical contamination and during medical evacuaCion.
The history of Che use of chemical weaponry shows convincingly that the
greatest damaRe is caused by OV when the army and population are noC pre-
par~d to combat the latsr consequences of a chemical attack. A high level
of preparedness in the population, all CD services and particularly the
medical service is a sure guarantee of keeping Cotal personnel losses to
a minimum, in particular by reducing the number of irretrievable losses and
changing the balance of injuries in favor of li~ht and semi-serious cases.
The material presented below has been arran~ed to correspond to the training
level of middle medical workers in problems of antichemical defense.
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.
run ~rri~~.e+L u~u u?vLi
[Text] '~able of ConCenCs
Cntroduction 3
ChapC~r 1 5horC descr~.ption of chemical weaponry 6
Ch~prer ~ Clinicgl meehods, diagnoais and tirearment o�
FP5 pnisotting 20
Chapeer 3 Clinical meChods, diagnosis and Creatment of
patients ~ffected by OV causing skin abscesses 69
ChapCer 4 Clinicnl meChods, diagnoeis and Creatment of
hydrocyanic acid poisoning 98
Chupter 5 Clinical meChods, diagnoeis and Creatment of
pnCienCs af�ected by psychomimetic OV 111
Chapter 6 Clinical methods, diagnosis and treatment of
patients affected by OV wiCh deteriorative effects 121
Ctiapter 7 Bnsic methods of resCoring respiraCory function 133 '
Chapr.er 8 Means of individual and collective antichemical
defense 150
I
Conclusion !
Liternture
~OPYRIGHT: Izdatel'stvo "Meditsina," Moscow, 1978
9354
CSO: 1840 END
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