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Genetic

Nadezhda Ivanovna Ryabokon, Igor Ivanovich Smolich and

Rose Iosiphovna Goncharova

Institute of Genetics and Cytology of National Academy of Sciences of Belarus, Minsk, Republic of Belarus

Keywords Population, Radiation, Genetics

Abstract Dynamics of population mutagenesis during 22 consecutive generations of animals,

as well as genetic radioadaptation were studied in natural populations of small mammals (bank voles) under chronic low-intensive irradiation due to the Chernobyl accident. The data obtained point to oppositely directed processes in irradiated populations: accumulation of mutations (genetic load of populations) and formation of genetic radioadaptation. It is suggested that the frequencies of genetic damages in populations could be higher in the absence of radioadaptation process. A relationship between the frequencies of cytogenetic injuries and low doses of radiation was revealed in animal generations studied. The non-linear dose-effect curves are most likely to be defined by the complicated microevolutionary processes in populations. The results obtained indicate the absence of genetic effect threshold of low dose radiation. Besides, they show that a dependence of cytogenetic effects on radiation low doses in series of irradiated generations cannot be revealed using linear equations.

Introduction

For predicting remote genetic consequences and working out the systems of population protection against negative effects of chronic ionizing radiation, it is necessary to know peculiarities of mutation process dynamics in a number of chronically irradiated animal generations, as well as quantitative ``dose-effect'' dependencies for different types of mutations in these generations.

In classical radiation genetics, there are experimental data on distinctive features of mutagenesis dynamics in chronically irradiated laboratory populations of Drosophila (Wallace, 1956) and unicellular algae (Shevchenko and Pomerantseva, 1985). Increased frequencies of mutations forming genetic load of populations remain in the populations studied during dozens of generations under long-term irradiation with high doses. Four stages were

The basic investigations represented in the given work were carried out within the framework of the State Programme of the Republic of Belarus for Minimizing and Overcoming Consequences of the Chernobyl Accident. Investigations of dynamics of the polyploid cell frequency in bank vole populations were partially supported by the J.D. and K.T. MacArthur Foundation (individual grant No. 95-31014 A-FSU).

The authors are grateful to Dr M.V. Malko (Institute of Physical and Chemical Radiation Problems, National Academy of Sciences of Belarus) for his consultations in estimation of absorbed doses, and to the staff of Antimutagenesis Laboratory (Institute of Genetics and Cytology, National Academy of Sciences of Belarus) for skilled technical help and helpful discussions.

The research register for this journal is available at

http://www.mcbup.com/research_registers/emh.asp

The current issue and full text archive of this journal is available at

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distinguished in dynamics of mutagenesis and genetic load: growth period (accumulation of mutations), period of stabilisation (equilibrium between induction of mutations and selection effects on them) and then reduction in the mutation frequency (increase in radioresistance) and stabilisation at a new lower level of mutagenesis (Wallace, 1956; Shevchenko and Pomerantseva, 1985; Dubinin, 1966). The UN Scientific Committee on the Effects of Atomic Radiation recommended the mutagenesis level at the equilibrium state (the second stage in dynamics of radiation mutagenesis) that was demonstrated in the experiments withDrosophilaand haploid algae (Wallace, 1956; Shevchenko and Pomerantseva, 1985; Dubinin, 1966) for estimating genetic risk of hereditary disease emergence in distant generations of chronically irradiated human population (United Nations, 1986; Shevchenko, 1996).

As for dynamics of the mutation process in chronically irradiated animal populations of higher taxonomic group, there were fragmentary data indicative of an increased level of mutagenesis in somatic and germ cells of rodents representing 20th-80th animal generations since the onset of additional chronic radiation impact (Shevchenko and Pomerantseva, 1985; Bashlykovaet al., 1987; Maslovaet al., 1984; Radioecology of Biocenosis, 1987; Grigorjev and Taskaev, 1985; Cristaldiet al., 1985, 1991; Taskaev, 1984; Gilevaet al., 1996; etc.).

Long-term monitoring of natural populations of animals living in radiocontaminated areas, in combination with a complex of radioecological and genetic methods of investigations, and appropriate approaches to statistically processed data is strongly required.

We have conducted such investigations in natural (free-living) populations of the European bank vole (Clethrionomys glareolus, Schreber) which is an indicator species of the environmental quality. The results obtained are presented in this paper.

Methodology

Methodology of the investigation is based on studying the dynamics of radiation loads and mutation process, as well as genetic radioadaptation in populations of bank vole living under chronic low-intensity irradiation after the Chernobyl accident during many generations.

Large forest areas for long-term monitoring were chosen in the territory of the Republic of Belarus at different distances from the Chernobyl NPP, with limited people activities and different radiocontamination densities of soil surface. The levels of137Cs contamination at monitoring sites (st.) 1, 2, 3 and 4 were 8, 18, 220 and 1,526kBq/m2 respectively. Summer-autumn trappings of murine rodents were carried out in 1986-1996: the first one ± in August-September 1986, i.e. in the four months following the accident (at the st. 2 ± in August 1991).

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completely consisted of no less than two new generations. Thus, from 1986 to

1996 we carried out monitoring work during approximately 22 irradiated generations of voles.

-irradiation dosage rates at monitoring sites were measured by dosimeters SRP-68-01-T and DBG-06T. The radiometric analysis of soil samples and whole-body frozen animals were mostly performed using -spectrometry ADCAM-300 (ORTEC, detector GEM-30185). Specific activity of 90Sr and transuranic elements was determined by radiochemical methods. Conventional formulas and coefficients were used when calculating absorbed doses (Moiseev and Ivanov, 1990). The individual absorbed doses were calculated in about 1,200 bank voles captured from four sites in 1986-1996.

The levels of mutation process were estimated using conventional methods: in somatic cells, metaphase analysis of bone marrow cells (Adler, 1984) and micronuclei assay of polychromatic (immature) erythrocytes (PCE) of bone marrow (Schmid, 1975; Adler, 1984) and normochromatic (mature) erythrocytes (NCE) of peripheral blood; in germ cells, abnormal sperm head (ASH) assay (Wyrobeket al., 1984). The total number of animals analysed exceeds 500.

Assessment of the embryonal mortality level before and after implantation was performed in 175 pregnant bank vole females by the conventional approaches also (Anderson, 1984).

A regression analysis was used to estimate peculiarities of radiation load dynamics and of mutation process dynamics, as well as to study relationships between individual dose loads and individual genetic effects.

To study genetic radioresistance of chronically irradiated populations, the captured animals were additionally exposed to acute 10, 50, 100 and 400 cGy dose of -rays from a 137Cs source at exposure dose rate 5.2R/min and the frequencies of micronucleated polychromatic erythrocytes were recorded at 24h after treatment.

Dynamics of radiation loads and mutation process in chronically irradiated populations

External and internal irradiation induced by caesium isotopes (137Cs, 134Cs) made a basic contribution to the total absorbed dose in the animals studied. Radiation load on the populations was the highest in the year of the accident and decreased greatly in the posterior ten years. Thus, the average rate of the total absorbed dose in voles at the st. 1, 3 and 4 decreased 2-15 times (Figure 1). ``Time-absorbed dose rate'' relationships were approximated by exponential functions.

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doses reached initially 1Gy only in some individuals at the st. 4. However, for first two years the radiation loads in animals at the st. 3 and 4 were underestimated because we did not have data on external‡-irradiation.

Using the metaphase method for counting the frequencies of chromosome aberrations (CA) and polyploidy in bone marrow cells, as well as micronucleus assay of PCE of bone marrow, it was revealed that an increased level of the somatic cell mutability remained during the whole period of observations in the animals in monitoring regions (st. 14). Thus, the CA frequencies were 3-7 times and polyploid cell (Pp-cell) ones were up to 300 times higher than the background (pre-accident) levels. The frequencies of micronucleated PCE (MN-PCE) reached double excess of background levels.

After the Chernobyl accident there were accumulated data on increased CA frequencies in somatic cells of the first generations of fish reared in radiocontaminated areas (Goncharova et al., 1996), in some generations of

Figure 1.

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murine rodents (Gecen, 1987; Eliseevaet al., 1996; Rakin and Bashlykova, 1996)

and amphibians (Eliseeva et al., 1996) inhabiting the Chernobyl radiocontaminated trace in 1986-1995.

Only not long ago was published information on remaining increased frequencies of cytogenetic injuries in somatic cells (peripheral blood) of the first to third generations of people in Altai region after long-term irradiation induced by explosions at Semipalatinsk testing ground (Suskov et al., 1997) and in the first to second generations of people living in the East-Uralian radioactive trace contaminated with 90Sr-90Y due to the accident in ``Mayak'' (Suskovet al., 1997).

However, the investigations (Shevchenko and Pomerantseva, 1985; Gecen, 1987; Eliseeva et al., 1996; Rakin and Bashlykova, 1996; Suskov et al., 1997; Suskov et al., 1997) in both pre- and post-Chernobyl periods had scanty material available or had no necessary statistical approaches to revealing peculiarities of the mutation process dynamics in chronically irradiated populations of animals and humans.

As a result of our monitoring, a large body of information on CA and polyploidy was amassed for estimating the dynamics of the mutation process in somatic cells over a series of bank vole generations. A regression analysis has revealed the relationship, common for the populations studied (st. 1, 3 and 4), between the mutagenesis level in somatic cells and the number of irradiated generations. The relationship for CA and Pp-cells was approximated by the exponential function and the second order parabola respectively (Figure 2).

Figure 2.

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This way, distinctive features of dynamics of radiation mutagenesis in populations of mammals were revealed. In particular, a gradual rise in the frequency of CA in somatic cells was observed up to the 22nd generation of animals since the onset of radiation impact. The number of Pp-cells increased in the 1st-12th generations and decreased by the 22nd one.

An increase in the frequencies of structural (CA) and genome (Pp-cells) mutations during many generations of animals at decreasing radiation doses points to higher sensitivity of somatic cell genomes of subsequent animal generations to low-intensive irradiation as against sensitivity of the previous generations. It could be caused by accumulation of genetic load in the populations and by rise in genomic instability.

Dynamics of the mutation process in somatic cells of a number of generations of chronically irradiated animals are in agreement with the above-mentioned literature data on dynamics of mutagenesis in germ cells of

Drosophila and in unicellular algae (Wallace, 1956; Shevchenko and Pomerantseva, 1985; Dubinin, 1966). However, the earlier revealed regularities in algae (Wallace, 1956; Shevchenko and Pomerantseva, 1985; Dubinin, 1966) are defined for the cases when the analysed mutations can be passed from one generation to another: in Drosophila, through germ cells; in algae, from maternal cell to daughter ones. We have studied the dynamics of somatic mutations emerging in each animal generation de novo. Nevertheless, accumulation of genetic load in germ cells of irradiated animals could give rise to instability of somatic cell genomes that result in increase in the frequencies of somatic mutations in consecutive generations of animals.

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Similar data were obtained by Pomerantseva et al. (1996). The authors

revealed that in 1987-1993 there was no additional increase in initially elevated frequencies of ASH, reciprocal translocations and recessive lethal mutations in wild house mouse populations living in areas contaminated with radionuclides after the Chernobyl disaster (maximal total absorbed doses in male gonads did not exceed 34Gy/month in 1986-1987 and gradually decreased). Pomerantseva

et al. assumed that radiation-induced mutations may lead to elimination of germ cells and of mice heterozygous with decreased viability. These processes result in removing excess mutations from populations. Only genetic damages that do not influence the viability of germ cells and early embryonal development stay in populations. And these mutations consist of genetic load of populations (Pomerantsevaet al., 1996).

We consider that statistically considerable rise in the frequency of embryonal losses up to the 22nd post-accident generation of bank vole is also caused by accumulation of genetic load in the populations studied (Figure 3). Some similarity between the dynamics of the relative quantity of embryonal lethals in females and that of the CA frequency in somatic cells of adult animals could be caused by the fact that most of embryonal lethals are due to CA in germ and zygotic cells. As the genetic load is accumulated in chronically irradiated populations, genome sensitivity of both somatic and germ cells to radiation increases that results in a rise in the CA frequency and then in embryonal mortality.

It should be noted that relationship between the dynamics of population density and that of embryonal lethality was not revealed. At the same time, very high frequencies of embryonal losses in monitoring populations of bank voles after the Chernobyl accident (Figure 3) point to radiation causality of the observed effects. These frequencies had a multiple excess over both the pre-accident levels in population at the st. 2 (Razhdestvenskaya, 1984) and the known background frequencies of embryonal lethals, typical for the whole habitat area of bank vole under different ecological conditions (Bashenina, 1981).

Significantly higher losses before implantation as compared with late embryonal lethality attract attention. They increased over the period of observations more than four times, being 30-50 times the pre-accident levels.

Figure 3.

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This is in agreement with the known data on high sensitivity of postovulatory ova and pre-implanted embryos to irradiation (United Nations, 1986; Mollset al., 1980; Nomura, 1984; etc.) and with the data on increased losses prior to implantation in root voles (Microtus oeconomus) inhabiting for above 20 years (approximately over 40 generations of rodents) uranium-radium contaminated sites at low radiation doses (about 2cGy/year) (Maslovaet al., 1974).

It was noted earlier that the moment of implantation is a functional test of the zygote state (Lyaginskaya and Smirnova, 1963). There could be underestimation of radiation effects for embryons when pre-implantation losses are not analysed. Subsequent stages of embryonal development proved to be more radioresistant (United Nations, 1986). Nevertheless, increased embryonal lethality after implantation was also observed in our populations of bank vole and populations of other species of murine rodents (Amvrosjev et al., 1993) at low doses of external and internal irradiation in the Chernobyl radioactive trace. Besides, there is information on greatly increased (up to 48 per cent) frequencies of embryonic malformations in bank and root voles representing approximately the 11th-18th post-accident generations of animals (1991-1994) inhabiting Bryansk Region (Russian Federation) radiocontaminated by Chernobyl fallout (Krylova, 1998). The high frequencies (by 40-83 per cent) of congenital developmental malformations were in children born in 1987-1995 in the regions of Belarus affected by the Chernobyl accident (Lazjuket al., 1998).

Main features of the mutation process dynamics revealed in chronically irradiated populations of small mammals within 22 animal generations could point to high genetic risk for existing and subsequent generations of humans and animals living in contaminated areas.

The data obtained can be used and were partially used (Goncharova, 1997; 1998) in predicting remote genetic consequences for human populations under chronic low dose irradiation. At the same time they display the necessity for long-term comprehensive and combined investigations of populations of humans and animals under chronic or prolonged low-intensive irradiation.

Formation of genetic radioadaptation in populations under low-intensive irradiation

The latter two stages in dynamics of radiation mutagenesis (reduction in the mutation frequency and stabilization at a new lower level of mutagenesis) that were revealed in laboratory populations of haploid algae (Shevchenko and Pomerantseva, 1985) andDrosophila(Wallace, 1956) define formation of genetic radioadaptation in populations. Adaptation of populations to mutagenic radiation impact is a long-term process. Thus, radioresistant clones of unicellular algae appeared in experimental populations after chronic irradiation of several tens of generations (Shevchenko and Pomerantseva, 1985). Radiosensitive individuals are gradually eliminated from the populations during that time and in the posterior period.

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algae (Shevchenko and Pomerantseva, 1985), higher plants (Shevchenko and

Pomerantseva, 1985; Dubinin et al., 1980), gastropods (Shevchenko and Pomerantseva, 1985; Dubinin et al., 1980) as well as of murine rodents (Shevchenko and Pomerantseva, 1985; Iljenko and Krapivko, 1989) inhabiting for some years the East-Uralian radioactive trace. Radioresistant individuals were shown to have more active repair systems. In the 25th-30th irradiated generations of murine rodents (northern redbacked vole ±Clethrionomys rutilis, and long-tailed field mouse ± Apodemus silvaticus) (Shevchenko and Pomerantseva, 1985) under additional administration of90Sr-90Y were revealed, on the one hand, increased total radiosensitivity (Shevchenko and Pomerantseva, 1985) and, on the other hand, high genetic radioresistance analysed for the frequency of CA in bone marrow cells (Shevchenko and Pomerantseva, 1985) and spleen (Iljenkoet al., 1980). The total radioresistance that was determined from the viability of animals following additional external

-radiation increased by the 30th-40th generations of rodents (Iljenko et al., 1989).

Based on these data, the process of population adaptation to chronic irradiation was noted to be very complicated and to proceed in different ways. This can be selection of radioresistant forms induced by chronic irradiation or existing earlier, as well as temporary activation of repair systems (Shevchenko and Pomerantseva, 1985; Iljenko and Krapivko, 1989).

Clearly defined features of the dynamics of Pp-cells in bank vole populations in Chernobyl radioactive trace (Figure 2) could indicate the processes of genetic adaptation in series of irradiated generations of animals.

We have compared genetic radioresistance of bank vole populations living in areas with different levels of radiation loads. Under additional acute -exposure of the 21st-22nd irradiated bank vole generations and application of micronucleus assay for PCE of bone marrow, the population inhabiting the st. 4 with high radiocontamination density was revealed to differ in higher radioresistance than rodents from the less contaminated st. 2 (Figure 4). So, the formation of genetic radioadaptation has begun by the 21st-22nd animal generations in the populations with higher radiation load.

Figure 4.

Cytogenic effects in bone marrow erythrocytes of bank voles after additional acute-irradiation from

137

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Thus, genetic radioadaptation was revealed to be formed rather earlier in murine rodent populations at our monitoring sites than it was shown in papers (Shevchenko and Pomerantseva, 1985; Iljenko et al., 1980), under conditions when absorbed doses were in the range from 1 to 100Gy (Gilevaet al., 1996).

Formation of genetic radioresistance in chronically irradiated populations is associated with the changes in functioning of complex system ``adaptive response'' and mechanisms of biological protection of tissues.

In particular, we have revealed functioning of system ``adaptive response'' in the 21st-22nd irradiated bank vole generations at the st. 4. Thus, under successive -irradiation of voles with an adapting 10cGy and then damaging (challenging) dose 100cGy the cytogenetic effect (frequencies of MN-PCE of bone marrow) was considerably lower than the effect of single irradiation with 100cGy. Besides, a sharp reduction in the frequencies of micronucleated mature (normochromatic) erythrocytes (MN-NCE) of peripheral blood in comparison with increased frequencies of immature erythrocytes with cytogenetic injuries (MN-PCE) in hematopoetic tissue (bone marrow) was revealed in the 21st-22nd vole generations (st. 14). This fact could be fully explained by elimination of cells with cytogenetic damages to protect peripheral blood of chronically irradiated animals against defective cells.

So, the pursued investigations have shown that the mutagenesis levels (frequencies of CA, Pp-cells and micronuclei) observed in the 21st-22nd generations of animals resulted from oppositely directed processes in irradiated populations: accumulation of mutations (genetic load of populations) and formation of genetic radioadaptation. The recorded frequencies of genetic damages in populations could be higher in the absence of radioadaptation.

However, it should be kept in mind that for adaptation, populations pay by elimination of the least adapted and the least viable individuals.

Dose-effect relationships for cytogenetic injuries in somatic cells of chronically irradiated animals

The relationship between individual frequencies of cytogenetic injuries (CA, Pp-cells and MN-PCE in bone marrow as well as MN-NCE in peripheral blood) and low levels of individual radiation loads in animals (st. 14) was revealed by using a regression analysis of the data. So, the relationship between the frequencies of cytogenetic injuries and concentration of incorporated radionuclides in the range of 4-145,410Bq/kg, dosage rate from 2 to 730Gy/ day and the total absorbed dose in the range of 0.02-7.3cGy was shown (Table I). The animals inhabiting the st. 1-4 were pooled in one sample within every year of investigations. In that way, analysed groups of animals represent individuals of approximately the same post-accident generations in the gradient of radiation loads (Table I, Figure 5).

It should be noted that causality of the observed cytogenetic injuries due to low radiation doses is followed over 22 generations of animals.

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overwhelming majority of the data (for separately considered populations or at their pooling) was better approximated by a polynomial function (Table I, Figure 5). The non-linear ``dose-effect'' relationships could be explained by peculiarities of low dose effects. But a different radiation history of populations and complicated microevolutionary processes in each irradiated population were most likely to increase population variability in individual radiosensitivity and in efficiency of biological system protection against

Table I.

1986 (1-2) 42 38-24,844 0.13* 6-670 0.13* 0.4-73 0.22*

1987 (3-4) 35-36

3,959-145,410

0.17* 205-615 0.17* 3-30 0.23**

1988 (5-6) 38-43 58-385,810 0.07 3-730 0.12* 0.2-267 0.03

1991 (11-12) 32-41 5-20,736 0.48** 3-132 0.23** 0.2-11 0.31**

1996 (21-22) 37 4-2,911 0.27** 2-46 0.15* 0.3-23 0.21**

Polyploid cells

1986 (1-2) 42 38-24,844 0.06 6-670 0.12* 0.4-73 0.16**

1987 (3-4) 35-36

3,959-145,410

0.16* 205-615 0.16* 3-30 0.02

1988 (5-6) 38-43 58-385,810 0.17** 3-730 0.12* 0.2-267 0.10

1991 (11-12) 32-41 5-20,736 0.32** 3-132 0.21** 0.2-11 0.41**

1996 (21-22) 37 4-2,911 0.06 2-46 0.11 0.3-23 0.06

Notes:R2Coefficient of determination; *p< 0.05 and **p< 0.01

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injuries (including repair systems). This could lead to complicated forms of dose-effect curves for cytogenetic injuries in somatic cells of a number of animal generations.

The results obtained point to the absence of genetic effect threshold of low doses of combined external and internal irradiation. Besides, they show that a dependence of genetic effects on radiation low doses in series of irradiated generations could not be revealed by using linear equations.

Concusions

Combined genetic and radioecological methods of investigation as well as application of a regression analysis for describing the mutation process dynamics and for estimating the relationship between individual frequencies of cytogenetic injuries and individual dose loads made it possible to obtain new knowledge on peculiarities of the genetic process dynamics in chronically irradiated natural populations of mammals and to determine the quantitative relationships between the frequencies of cytogenetic injuries and low doses of irradiation.

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Shevchenko, V.A. (1996), ``Assessment of genetic risk of human population irradiation'', in Burlakova, E.B. (Ed.),Consequences of Chernobyl Catastrophe: Human Health(in Russian), Centre of Ecological Politics of Russia, Moscow, pp. 50-67.

Shevchenko, V.A. and Pomerantseva, M.D. (1985),Genetic Consequences of Ionizing Irradiation

(in Russian), Nauka, Moscow.

Suskov, I.I., Dubrovina, T.B., Safonova, N.V. and Shevchenko, V.A. (1997), ``Family analysis of aberrant chromosomes in rural populations'', 3rd Congress on Radiation Research. Radiobiology, Radioecology, Radiation Safety: Abstracts, Moscow, 14-17 October, International Association of Academies of Sciences, Pushchino(in Russian) Vol. 2 (section IV-VI), pp. 84-5.

Suskov, I.I., Elisova, T.V., Dubrovina, T.V.et al.(1997), ``Family analysis of remote genetic effects of chronic radiation impact in habitants of village Muslyumovo'', 3rd Congress on Radiation Research. Radiobiology, Radioecology, Radiation Safety: Abstracts, Moscow, 14-17 October, International Association of Academies of Sciences, Pushchino(in Russian), Vol. 2 (section IV-I), pp. 85-6.

Taskaev, A.I. (Ed.-in-chief) (1984),Radiation as Ecological Factor under Antropogenic Pollution: Transactions of Komi branch, Academy of Sciences of USSR (in Russian), No. 67. Syktyvkar.

United Nations (1986),Genetic and Somatic Effects of Ionizing Radiation. Report of the United Nations Scientific Committee on the Effects of Atomic Radiation to the General Assembly with Annexes, United Nations Publication E.86.IX, United Nations, New York, NY. Wallace, B. (1956), ``Studies on irradiated populations ofDrosophila melanogaster'',Journal of

Genetics, Vol. 54 No. 2, p. 280.

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[S e a r ch] [Ad v a n ce d S e a r ch] [S u b m it Re s e a r ch] [Ed it Re s e a r ch] [He lp]

We lco m e t o t h e En v ir o n m e n t a l Ma n a g e m e n t & He a lt h In t e r n e t Re s e a r ch Re g is t e r , d e liv e r e d b y MCB Un iv e r s it y Pr e s s in a s s o cia t io n w it h t h e I n t e r n a t io n a l Jo u r n a l o f S u s t a in a b ilit y in Hig h e r

Ed u c a t io n a n d t h e fo llo w in g in t e r n a t io n a l b o d ie s :

-THE INTERNATIONAL FEDERATION OF ENVIRONMENTAL HEALTH

Caring for the Environment in the Interests of World Health

Co - s p o n s o rs o f t h e En v iro n m e n t a l Ma n a g e m e n t & He a lt h I n t e rn e t Re s e a rc h Re g is t e r

Th e "En v ir o n m e n t a l Ma n a g e m e n t a n d He a lt h " ( EMH) In t e r n e t Re s e a r ch Re g is t e r r e p o r t s o n cu r r e n t r e s e a r ch w o r ld w id e , in clu d in g s t u d ie s co m p le t e d d u r in g t h e p a s t s ix t o n in e m o n t h s . Re s e a r ch m a y b e lin k e d t o a n a ca d e m ic co u r s e o f s t u d y o r p r o je ct , b e in d e p e n d e n t , o r

o r g a n is a t io n - b a s e d . Bo t h in d u s t r y a n d t h e a ca d e m ic w o r ld a r e r e p r e s e n t e d , a s w e ll a s t h e g o v e r n m e n t a n d NGO s e ct o r s . Th e r e g is t e r a im s t o b e co m p r e h e n s iv e in it s g e o g r a p h ic a n d

Sponsors

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s u b je ct co v e r a g e a n d a im s t o m a in t a in a s o u n d b a la n ce b e t w e e n t h e o r y a n d p r a ct ice . En t r ie s w h ich r e p o r t o n a p p lie d r e s e a r ch a r e p a r t icu la r ly w e lco m e .

Th e Re g is t e r in clu d e s , b u t is n o t r e s t r ict e d , t o r e s e a r ch in t h e fo llo w in g a r e a s o f e n v ir o n m e n t a l m a n a g e m e n t a n d e n v ir o n m e n t a l h e a lt h : g e n e ra l e n v iro n m e n t a l m a n a g e m e n t ; e n v iro n m e n t a l

p o licie s a n d le g is la t io n ; in d u s t ria l e n v iro n m e n t a l p ra ct ice s a n d p ro ce s s e s ; e n v iro n m e n t a l h e a lt h ; a p p lie d t ra in in g in e n v iro n m e n t a l m a n a g e m e n t ; e n v iro n m e n t a l m a n a g e m e n t s y s t e m s ; s u s t a in a b le d e v e lo p m e n t , la n d u s e a n d p la n n in g ; w a s t e s m a n a g e m e n t ; e n e rg y a n d it s u s e ; w a t e r, w a t e r u s e , w a s t e w a t e r a n d w a t e r m a n a g e m e n t ; d is e a s e s a n d illn e s s e s d e riv in g fro m e n v iro n m e n t a l

p ro b le m s ; a s p e ct s o f e n v iro n m e n t a l m a n a g e m e n t a n d la w .

Th e b e n e fit s o f re g is t e rin g y o u r re s e a rc h : Th e EMH In t e r n e t Re s e a r ch Re g is t e r p r o v id e s t h e

r e s e a r ch co m m u n it y w it h p r e - p u b lica t io n in fo r m a t io n a n d t h e p o t e n t ia l fo r fu r t h e r n e t w o r k in g , a lo n g s id e e a r ly in fo r m a t io n o n n e w a r e a s fo r a p p lica t io n a n d d e v e lo p m e n t in t h e fie ld . It a ls o a llo w s t h e r e s e a r ch e r t o :

* p r o m o t e t h e ir o w n r e s e a r ch a n d t h a t o f t h e ir o r g a n is a t io n / in s t it u t io n ; * e n s u r e t h a t t h e y a r e n o t d u p lica t in g r e s e a r ch t h a t is a lr e a d y u n d e r w a y ; * id e n t ify p o s s ib le r e s e a r ch m e t h o d o lo g ie s ;

* id e n t ify p e e r s fo r co lla b o r a t iv e r e s e a r ch p r o je ct s ; * id e n t ify p o s s ib le s o u r ce s o f fu n d in g fo r r e s e a r ch ;

* id e n t ify t y p e s o f r e s e a r ch u n d e r w a y , e . g . t h e o r e t ica l, a p p lie d r e s e a r ch , ca s e s t u d y ; * id e n t ify a r e a s w h e r e fu r t h e r r e s e a r ch is r e q u ir e d .

Th e EMH In t e r n e t Re s e a r ch Re g is t e r is fr e e ly a v a ila b le t o a ll w h o r e g is t e r t h e ir r e s e a r ch , t o s u b s cr ib e r s t o t h e a b o v e jo u r n a ls a n d t o m e m b e r s o f a s s o cia t e d o r g a n is a t io n s / in s t it u t e s . Ple a s e co n t a ct Je n n y Pick le s : jp ick le s @m cb . co . u k fo r d e t a ils o f h o w y o u r a s s o cia t io n ca n g a in fr e e a cce s s .

All e n t r ie s a r e v a lid a t e d b y Pr o f. Wa lt e r Le a l Filh o

,

Te ch n ica l Un iv e r s it y Ha m b u r g - Ha r b u r g

Te ch n o lo g y Tr a n s fe r ( TUHH/ Tu Te ch ) , Ge r m a n y a n d t h e Ro y a l In s t it u t e o f Te ch n o lo g y , S t o ck h o lm , S w e d e n .

No r e s p o n s ib ilit y is a cce p t e d fo r t h e a ccu r a cy o f in fo r m a t io n co n t a in e d in t h e r e s e a r ch p r e s e n t e d w it h in t h is Re g is t e r . Th e o p in io n s e x p r e s s e d h e r e in a r e n o t n e ce s s a r ily t h o s e o f t h e Ed it o r s o r t h e p u b lis h e r .

© MCB Un iv e rs it y Pre s s , UK a n d Th o m a s Te ch n o lo g y S o lu t io n s ( UK) Lt d .

[S e a r ch] [Ad v a n ce d S e a r ch] [S u b m it Re s e a r ch] [Ed it Re s e a r ch] [He lp]

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