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P"�yN]N]N�y�#�i�-4��4^hT5L[5�-4-4-4�q�q,S<-4-4-4�s�6�6�6�6���������������������������������������������������������������������y-4-4-4-4-4-4-4-4-4L(t":"XENOBIOTICS  -450  DEPENDENT METABOLISM IN PATHOGENESIS OF MALE REPRODUCTIVE DISORDERS
Ganna M. Shayakhmetova, Larysa B. Bondarenko 

ABSTRACT
Last decades investigations demonstrated significant decreasing of men fertility and sperm quality. Approximately 50% clinically confirmed infertility cases are attributed to the male partners. Male infertility pathogenesis is investigated insufficiently; its direct interrelation with such pathologies as varicocele or urogenital infections was demonstrated only in 23% of cases. At present it is supposed that different etiologic factors (including idiopathic) could start similar processes which final result would be spermatozoids quality and/or quantity decreasing. The present review confirms possibility of cytochrome P-450 (CYP2E1) isoforms activities modulation directly in male reproductive organs, what could be one of the mechanisms of subfertility development and could mediate paternal negative effects on the posterity. Current scientific data specify great importance of multiple cytochromes P-450 isoforms investigations in extra-hepatic tissues. And among them CYP2E1 is of special interest, as CYP2E1-mediated metabolism of xenobiotics directly in male reproductive organs could produce toxic intermediates, which are able to interact with vitally important cells structures, and oxygen reactive species overproduction with the further development of oxidative stress. From our point of view for exact detection of all events and mechanisms, involved into development of testicular toxicity, the evaluation of cytochrome P-450 isoforms inhibitors and inducers combined action in vitro and in vivo could be very perspective. Such investigations allow working out effective methods for preventing men�s infertility under increasing pressure of adverse environmental factors, especially chemicals.

Key words: male reproductive disorders, testes, cytochrome -450, xenobiotics.

Authors information:
Ganna M. Shayakhmetova & Larysa B. Bondarenko 
General Toxicology Department,
SI  Institute of Pharmacology & Toxicology of National Academy of Medical Sciences of Ukraine�, 
Eugene Pottier 14, Kyiv, 03680, Ukraine,
Tel: +380 44�456 78 65
Fax:  +380 44�456 63 44
E-mail:  HYPERLINK "mailto:anna_shaaykhmetova@yahoo.com"anna_shayakhmetova@yahoo.com;  HYPERLINK "mailto:larabon04@yahoo.com"larabon04@yahoo.com

Last decades investigations demonstrated significant decreasing of men fertility and sperm quality [1-4]. Approximately 50% clinically confirmed infertility cases are attributed to the male partners [5, 6]. 
Normal male reproductive capacity is a result of complex interactions of numerous mechanisms involved into spermatozoids production and their transport from gonads to ejaculate.  Hypothalamohypophysial system regulates functioning of Sertoli cells, spermatogenic epithelium and androgens production by Leydig cells via producing gonadotropin, follicle-stimulating hormone and luteinizing hormone. Functioning of cells, localized outside of seminiferous tubules (Leydig cells and myoepithelial cells), provides adequate level of spermatozoids production. The integrity of blood-testis barrier is essential for meiosis unique microenvironment creation and post-meiotic germ cells development as it provides the isolation of these events from the systemic circulation, which would otherwise develop anti-sperm antibodies [7]. The accessory sex glands (epididymis, prostate and seminal vesicles) also play a role in realization of the spermatozoids fertilizing potential [8].  
Male infertility pathogenesis is investigated insufficiently; its direct interrelation with such pathologies as varicocele or urogenital infections was demonstrated only in 23% of cases [7]. At present it is supposed that different etiologic factors (including idiopathic) could start similar processes which final result would be spermatozoids quality and/or quantity decreasing [9]. 
Human male reproductive function could be injured by exposing to different xenobiotics-reproductive toxicants from environment, via occupational exposure, as a result of life-style (smoking) or chemotherapy [10-12]. Many different mechanisms are involved into the development of negative effects on human male reproductive function: 1) normal processes of endocrine or paracrine regulation disruption by hormone- or anti hormone-mimicking compounds; 2) direct or metabolically mediated destructive action on specific cell types, such as germ cells, Sertoli cells or Leydig cells; 3) blood-testis barrier damage; 4) gonads and liver enzymes systems induction or inhibition, which could cause increase or decrease of steroid hormones secretion or clearance; 5) mutagenic effects on germ cells, which could cause production of spermatozoids with low fertilizing potential or unable to fertilize ovum, developmental defects or posterity genetic diseases; 6) neurotoxic action, accompanied by erection/emission/ejaculation disorders [7, 13, 14]. It must be stressed that reproductive toxicants harmful actions could be realized by more then one mechanism.  For example, the halogenated polycyclic hydrocarbons may act indirectly by induction of microsomal monooxygenases or transferases. These compounds may also act directly by binding to steroid hormones receptors. Two indirect mechanisms are involved into development of polycyclic aromatic hydrocarbons reproductive toxicity � their metabolism to chemically reactive diol-epoxides and induction of microsomal  -450-dependent monooxygenases in liver and other organs [13].
The majority of xenobiotics require metabolic activation in order to exert adverse effects via covalent interactions between intermediate metabolites and cellular macromolecules, such as DNA or protein. Additionally, xenobiotics may gain properties of endocrine disruptors in processes of toxic bioactivation [15]. 
Great number of chemical compounds in mammals  organisms is metabolically activated by cytochrome  -450. In these enzymes  system investigators usually differentiate two great classes   cytochromes  -450 (induced by phenobarbital) and cytochromes  -448 (induced by 3-methylcholantrene) [16].  
Lake et al. studied these isozymes activities in testis supernatant (obtained at 10.000g) using biphenyl as substrate. Due to studies of 4-hydroxy biphenyl synthesis processes, activation of cytochrome  -450 was demonstrated, while cytochrome  -448 remained inactive [17].  Murray et al. using immunocytochemical method with monoclonal antibodies proved cytochrome  -450 presence in human testis Leydig cells [18]. Subsequently tissue distribution of mRNA expression of human cytochrome P-450 isoforms was assessed by high-sensivity real-time reverse transcription PCR. It has been shown that in human testes 31 isoforms were expressed: CYP1A1, CYP1A2, CYP1B1, CYP2A6/7,  CYP2A7, CYP2B6, CYP2C8, CYP2C9, CYP2C19, CYP2D6,  CYP2E1, CYP2F1, CYP2J2, CYP3A3/4, CYP3A4, CYP4A11, CYP4B1, CYP4F2, CYP4F3, CYP4F8, CYP5A1, CYP7B1, CYP8A1, CYP8B1, CYP11A, CYP17, CYP19, CYP27B1, CYP39A1, CYP46, CYP51 [19]. 
Testes cytochrome  -450 isoforms composition varies widely between species. For example in the Cynomolgus Monkey (Macaca fascicularis) testes mRNA expression of seven isoforms (!YP1A1, CYP1A2, CYP2D17, CYP2E1, CYP2J2, CYP3A5, CYP3A8) has been demonstrated [20]. 
Cytochrome  -450 isoforms mRNA expression in different tissues of frequently used experimental animals (rats and mice) also was studied. It has been shown, that only three isoforms of cytochromes subfamily 1 (CYP11, CYP12, CYP11) and one isoform of cytochromes subfamily 2 (CYP21) were present both in humans and in above mentioned animal species. They also had some interspecies specificity caused by differences in the amino acid sequences. For example, striking species differences were noted, as mouse CYP1B1 did not activate phenanthrene at all, while human CYP1B1 exhibited its significant metabolic turnover comparable to CYP1A1 and CYP1A2  [21]. 
Notably, that CYP1B1 protein is expressed at a relatively high level in rat testis (57 pmol/mg). It may be the most abundant P450 enzyme in testicular microsomes from adult rats. The reason why CYP1B1 is expressed at a high level in these tissues is not known [22].  
It must be stressed, that the testis contains two distinct sets of cytochrome P450 (P450) enzymes: first � steroidogenic P450 enzymes (e.g., CYP11A1 and CYP17A1), which catalyze key reactions in androgen biosynthesis [23], and the other set comprises those P450 enzymes, which could play a role in the biotransformation of hydrophobic xenobiotic compounds to more water-soluble metabolites [24]. The physiological role of xenobiotic-metabolizing P450 enzymes in testis is not definitively known. It has been supposed that they catalyze the oxidative biotransformation of lipophilic xenobiotic or endogenous compounds within the testes, what could be important for testicular toxicity realization [25]
The greatest impact into xenobiotics metabolism, including medical drugs, belongs to isoforms CYP1A2, CYP2C9, CYP2C19, CYP2D6, CYP2E1, CYP3A4 [26]. 
Only for small number of cytochrome P450 isoforms their involving into pathogenesis of male reproductive system disorders has been demonstrated at present. In particular, recent results demonstrating differential CYP3A4 expression in prostatic tissues are of great interest. They confirm that decreased CYP3A4 expression may contribute to the development of prostate cancer [27]. For certain combined CYP3A4/CYP3A5 haplotypes also was demonstrated differential susceptibility to this disease [28]. Recently discovered isoform CYP3A43 demonstrated low expression rate in liver, but simultaneously it expressed in prostate and testis. Its substrate specificity is currently unclear [28].  
Experiments on animals demonstrated that, due to their CYP-inducing potential, compounds with different chemical structure   mirex (inducer of CYP1A2, 210, 21 and 3), 2,2-bis(?-chlorophenyl)-1-1- dichloroethylene (inducer of CYP1A2 and 210), vinclozolin and flutamide (inducers of CYP3A and 12) may significantly influence on testosterone metabolism. Such effect may be a contributing factor in their antiandrogenic effects [29].
Among cytochrome P450 isoforms, which are expressing in male gonads, CYP21 is important from a toxicological point of view, taking into account its high inducibility and ability to massive generation of oxygen reactive species, such as, super oxide radical [30]. At present it has been demonstrated, that CYP21 participates in metabolism of wide spectrum of xenobiotics: alcohols, aldehydes, ketones, aromatic compounds, esters, fatty acids, halogenated and non halogenated alkanes and alkenes, nitrosamines, azocompounds. This list includes solvents widespread in industry and everyday life, medicines, endogen compounds, etc. [30]. Mainly investigators� attention is spared to CYP21 inducers effects on liver, although xenobiotics oxidative metabolism could take place not only in this organ. Now cytochrome  -450 21 presence has been demonstrated in human kidney, adrenal glands, lung, brain, testis, prostate, uterus, placenta, nasal mucosa [19].  
At 1998 Jiang et al. tried to identify mRNA of cytochrome  -450 21 in rat testis and prostate using methods of reversed transcriptase polymerase chain reaction, Southern-blotting and DNA sequencing [31]. Immunoblot analysis has allowed detection of CYP21 protein in testis, but not in prostate. Similar results authors obtained for activity of CYP21 marker enzyme  p-nitrophenolhydroxylase [31]. Exposure of rats to typical cytochrome  -450 21 inducer   pyridine resulted in a 2.9-fold increase of p-nitrophenol hydroxylation by testicular microsomes. At the same time diethyldithiocarbamate, a selective mechanism-based inhibitor of P450 2E1, or a P450 2E1 monoclonal antibodies, caused marked inhibition of testicular microsomal p-nitrophenol hydroxylase activity. Thus, these studies clearly demonstrated that inducible cytochrome  -450 21 was present in testis. This fact allowed authors to suppose this isoform possible role in the bioactivation of environmental chemicals to genotoxic metabolites directly in male gonads [31]. 
Later DuTeaux et al. [32, 33] and Forkert et al. [34] demonstrated CYP2E1 presence in rat seminiferous tubules, Leydig cells and epididymis.  Authors supposed, that these tissues would also participate in xenobiotics oxidative metabolism [32, 33, 34]. This fact is of great importance taking into account epididymis and seminiferous tubules great role in male reproductive function realization. 
Seminiferous tubules connect testicles with epididymis and absorb more than 75 % of liquid fraction during sperm transmission to epididymis [35, 36]. During migration through epididymis ductules, membranes of spermatozoids are modified, which is important to obtain motility and ability for oocyte impregnation. Germ cells, extracted directly from seminiferous tubules, are immobile and cannot penetrate oocyte membrane. Thus, final maturation of spermatozoids takes place in epididymis ductules and importance of this stage for males fertility is not subject a doubt [36-38]. And therefore xenobiotics toxic bioactivation in seminiferous tubules and epididymis would undoubtedly have negative effect on male reproductive function.
Role of CYP2E1 was proved in bioactivation of such testicular toxicants as acetone and benzene. Administration of benzene to mice caused testicular atrophy; spermatozoids number decreasing and increased number of their anomalous forms [39]. Acetones ability to influence on mice sperm quality (depressed sperm motility, cauda epididymal and epididymal weight decrease and elevated incidence of abnormal sperm) have been demonstrated [40]. This could be evidence that inducers of CYP2E1 could cause damage of reproductive organs and increase toxicity of potential testicular toxicant due to their metabolism modulation directly in male gonads. 
Effects of trichloroethylene (TCE) on male reproduction and fertility have been studied in mice and rats, and also assessed in workers exposed to TCE. CYP2E1 induction and oxidative metabolism appear to be important in the systemic toxicity and may likewise play a role in the reproductive toxicity of this compound [41].
I vivo study using male Sprague-Dawley rats exposed to TCE in drinking water at 0.2% or 0.4% (v/v) for 14 days was conducted. There were no treatment-related changes in testes and epididymis weight, sperm concentration, or sperm motility, but TCE caused slight histologic changes in efferent ductal epithelium, coinciding with ductal localization of CYP2E1. There were no alterations in the testis or in any segment of the epididymis, but the rats exhibited significant dose-related reductions in percent fertilized ova from untreated females in vitro [32]. Among this dose-dependent increase of lipoperoxydation was demonstrated in rat sperm, which could be related with expression of abovementioned cytochrome  -450 isoform. Sperm oxidative modification in its turn could cause males fertility loss [32]. In fact lowering of epididymis cells antioxidant abilities as a result of increased levels of oxygen reactive forms stipulates apoptotic changes in gametes and could be one of the main factors, limiting their fertility [42]. 
Immunoreactive CYP2E1 was localized to the efferent ducts and corpus epididymal epithelia. TCE metabolism was inhibited 77% when efferent duct microsomes were preincubated with antibodies to CYP2E1. This isoform inhibition by antibodies was the most expressed in epididymis (in comparison with testis). Dichloroacetyl adducts were detected in epididymal and efferent ducts microsomes exposed to TCE in vitro [33]. 
In experiments on mice (exposed to TCE by inhalation) immunoblotting confirmed that CYP2E1 protein was present in greater amounts in their epididymis than in testis. P-Nitrophenol hydroxylation, a CYP2E1 catalytic activity marker, was also higher in the epididymis than in the testis. Chloral, a major TCE metabolite, was generated in microsomal incubations at significantly higher levels in epididymis than in testis. Antibody inhibition of CYP2E1 reduced chloral formation, which was more pronounced in epididymis than in testis [34].
The authors suggested that CYP2E1-dependent metabolism of TCE to reactive metabolites and their subsequent covalent binding to cellular proteins may be involved in the male reproductive toxicity of trichloroethylene [32, 33, 34]. 
Forkert et al. examined eight mechanics with clinical infertility that had occupational exposure to TCE for at least 2 years. Seminal fluid from all eight subjects contained TCE, chloral, and trichloroethanol, whereas dichloroacetic acid and trichloroacetic acid were present in only two and one sample, respectively. Neither TCE nor its metabolites was detected in the five control male seminal fluid samples. CYP2E1 was found in normal human testes and epididymides, specifically in the Leydig cells in the intersticium of the testis and the caput (head), corpus (body), and cauda (tail) of the epididymis. This finding was consistent with findings in rodents, which demonstrated that CYP2E1 was involved in TCE metabolism in these tissues [43]. 
The contribution of CYP2E1 to 1-bromopropane (1-BrP) metabolism and the induction of male reproductive toxicity was investigated using wild-type (WT) and Cyp2e1-/- mice. In conditions of inhalation exposure the elimination half-life of [1,2,3-(13)C]-1-BrP was longer in Cyp2e1-/- mice relative to WT. At the same time in WT mice urine 1-BrP the main metabolite was mercapturic acid of 1-bromo-2-hydroxypropane, while in !C@251 -/- animals   products of conjugation of 1-BrP with glutathione. Despite a 170% increase in 1-BrP exposure in Cyp2e1-/- vs. WT mice, sperm motility in exposed Cyp2e1-/- mice did not change relative to unexposed matched controls. Both 1-BrP and 1-bromo-2-hydroxypropane inhibited the motility of sperm obtained from WT mice in vitro. However, only 1-bromo-2-hydroxypropane reduced the motility of sperm obtained from Cyp2e1-/- mice in vitro, suggesting that conversion of parent compound to 1-bromo-2-hydroxypropane within the spermatozoa may contribute, at least in part, to reduced motility. The author suggested that metabolism of 1-BrP is mediated in part by CYP2E1, and activation of 1-BrP via this enzyme may contribute to the male reproductive toxicity of this chemical [44]. 
It is well known that ethanol is able to inhibit testosterone production and cause testicular dystrophy [45]. According to one of the hypothesis, this phenomenon is connected with metabolic transformation of alcohol into acetaldehyde directly in testis. Investigation of ethanol oxidation to acetaldehyde in testis microsomes fraction demonstrated its enzymatic character and dependence on NADPH and oxygen presence. It has been shown that preparations of testis microsomes in the presence of NADPH were also able to produce both hydroxyl- and 1-hydroxyethyl- free radicals. Their generation was modulated by the presence of compounds, which inhibited acetaldehyde production � diphenyleneiodonium, gossypol, and deferoxamine. Such results evidenced the presence in testicular microsomal fraction of an enzymatic NADPH and oxygen-dependent ability to transform ethanol to acetaldehyde. The authors postulated at less the partial involvement of CYP2E1 and P450 reductase into the microsomal transformation of ethanol into acetaldehyde. Also other enzymes with lipoxygenase-/-peroxydase-like behavior could be involved [46]. 
It should be noted, that strong evidence for a microsomal ethanol oxidizing system in rat Leydig cells was provided [47]. Activity of the microsomal ethanol oxidizing system in Leydig cells was 47.4 +/- 4.1 nmol acetaldehyde per 20 min per mg protein, while activity in crude interstitial cells was 26.0 +/- 5.4 nmol. This allowed authors to suggest that among cells comprising interstitial cells, activity is concentrated in Leydig cells. Activity had linear dependence on protein concentration and incubation time. The most effective cofactor was NADPH. Inhibitors of alcohol dehydrogenase and catalase activities (4-methylpirazol and potassium cyanide) had no effects on alcohol oxidation by Leydig cells microsomes. Microsomal ethanol-oxidizing system activity here was twice higher, than at interstitial cells [47]. Taking into account ability of CYP2E1 to generate oxygen reactive species and their quick interaction with organic molecules, producing secondary free radicals and oxygen reactive species [30], such cascade could cause oxidative damage of Leydig cells. On the other hand Leydig cells damage could cause inhibition of testosterone production and this, in its turn could affect Sertoli cells function and negatively influence spermatogenesis [14].
Our recent investigations have been demonstrated that chronic ethanol consumption caused testicular failure along with an overexpression of CYP2E1 mRNA and protein in the testes as well as quantitative changes in type I collagen amino acid contents. The profound alcohol-mediated changes in collagen type I amino acid contents may have affected the spermatogenic epithelium state. The modulation of testicular cytochrome P450 2E1 mRNA and protein expression could change the functioning of this isozyme in target organs and take part in the mechanism of ethanol gonadotoxicity [48].
It is well known, that cytochrome  -450 21 participates in metabolism of inhalation anesthetics   halothane, isoflurane, sevoflurane, enflurane and desflurane to highly toxic metabolites (trifluoracetic acid, trifluoroacetyl chloride etc.), which acetylate proteins and cause neoantigenes formation [49, 50]. 
The induction of CYP2E1 in rats testes and liver and the presence of trifluoroacetylated (TFA) adducts in spermatozoa, testes, liver and plasma were demonstrated after subchronic exposition of halothane. After halothane exposure, p-nitrophenol hydroxylase activity increased 3.2-fold and CYP2E1 apo-protein content � 7-fold in testes, whereas in liver p-nitrophenol hydroxylase activity increased 2.3-fold and CYP2E1 apoprotein content � 1.4-fold. These results suggest a differential inductive effect of halothane on CYP2E1 in these tissues. Moreover, TFA adducts were present in microsomes of testis and liver and in plasma of halothane-treated rats. The immunoblot analysis of testicular microsomes showed two intense TFA protein bands of 63 and 59 kDa.  In addition, TFA adducts were detected by immunofluorescence in spermatozoa, probably in the acrosome and/or perinuclear theca region, and in the distal tail of spermatozoa. The authors suggest that halothane induces its own biotransformation both hepatically and extrahepatically and in addition, that the nature of the TFA adducts will depend on the proteins present in each tissue. Also, the presence of TFA adducts in spermatozoa may result from the activation of halothane in the reproductive tract [51]. 
It was shown that anesthesiologists� wives, not exposed to the operating room environment, presented an abortion rate 3 times higher and a premature birth rate 4 times higher than controls [52].  The study compared physicians exposed to anesthetics to non-exposed individuals, aiming investigation of chromosomal changes, has been performed. Cytogenetic analysis has shown an increased frequency in chromosomal aberrations per cell in the exposed group as compared to controls, confirming the clastogenic effect of such compounds [52].  CYP2E1 presence at male gonads and its inducibility from our point of view could be of great importance for genotoxic metabolites production during chemicals biotransformation. It could be supposed CYP2E1 involving into development of remote effects on the posterity of male, which were chronically exposed to inhalation anesthetics.
CYP21 is thought to play a significant role in the bioactivation of 1,3-butadiene (BD) to DNA reactive epoxide metabolites that induce somatic and germ cell genotoxicity in mice [53]. The presence of CYP2E1 in testes has been demonstrated by reverse transcriptase-polymerase chain reaction, immunoprecipitation-Western blotting and immunohistochemistry of tissue sections. Detection of CYP2E1 in the testes was limited to interstitial cells. The authors suggest that in situ bioactivation of BD in testes by CYP2E1 may contribute to BD-induced germ cell genotoxicity [53]. 
Acrylamide (AM) is commonly used in the synthesis of plastics and polymers. In rodents, AM is metabolized to the epoxides glycidamide (GA). The role of cytochrome P450 in the metabolism of AM has been determined. On the basis of evaluation of urinary metabolites, obtained data suggest that CYP2E1 is possibly the single cytochrome P450 enzyme involved into the metabolism of AM in mice [54]. CYP2E1-mediated metabolism of AM to GA, was shown to be a prerequisite step in AM-induced germ cell mutagenicity. Using Cyp2e1-null (Cyp2e1"/") and wild-type (Cyp2e1+/+) male mice, studies were performed in order to investigate the hypothesis that the formation of GA was responsible for the germ cell mutagenicity attributed to AA exposure. Untreated female mice mated with AM-treated Cyp2e1+/+ males demonstrated an increase in the number of chromosomally aberrant embryos (resorptions) and a subsequent decrease in the proportion of live fetuses, number of implants, and rate of pregnancy. In contrast, female mice mated to AM-treated Cyp2e1"/" males exhibited no significant adverse effects, indicating that AA-induced germ cell mutagenicity was dependent upon CYP2E1-mediated metabolism of AM [55]. Further investigations demonstrated that AM toxic effects on mice germ cells increased in obese male mice (pathological state, which is accompanied with CYP2E1 induction). On the base of this findings the authors suggest that obese men may be more susceptible to the effects of reproductive toxicants and possibly other environmental pollutants and carcinogens, especially those chemicals metabolized by CYP2E1 [56]. 
Taking into account all above mentioned, it is possible to assert that the earliest manifestations of remote consequences of chemicals, metabolizing via CYP2E1 system, for male reproductive system could have general character. It must be specially distinguished, that vitally important organs cell structures destabilization and thus damage of detoxification systems (liver, kidney), non-specific defense (macrophages), regulation (central nervous system) and reproducing (gonads, placenta) take place even then, when there is no any �obvious� structure-functional changes in organism. At the same time cell membranes destabilization remains in the future and in some cases even progresses with development of substantial disorders in reproduction, embryogenesis, non-specific resistance, behavioral reactions, mutagenesis, and carcinogenesis.
The present review confirms possibility of cytochrome P-450 isoforms activities modulation directly in male reproductive organs, what could be one of the mechanisms of subfertility development and could mediate paternal negative effects on the posterity. Now such effects have been investigated only for small number of compounds, but further their list can be extended.
The analysis of abovementioned findings suggests an idea that testicular toxicity of some xenobiotics metabolizing via cytochrome P-450 system is possibly based on general processes. This fact could be of great importance as male reproductive system is one of the main targets for great number of chemicals [7]. It is known that global environmental pollution significantly influences on male germ cells [15, 57]. Despite of the fact that testicular toxicants actions could be directed on different structures of reproductive system, final general answer would be, as a rule non-specific, as damage of one cells type causes whole cascade of events, mediating structural changes and dysfunctions of other parts of the system [14]. Current scientific data specify great importance of multiple cytochromes P-450 isoforms investigations in extra-hepatic tissues. And among them CYP2E1 is of special interest, as CYP2E1-mediated metabolism of xenobiotics directly in male reproductive organs could produce toxic intermediates, which are able to interact with vitally important cells structures, and oxygen reactive species overproduction with the further development of oxidative stress. 
It is necessary to notice, that great number of investigations stresses importance of finding out combined effects of xenobiotics on male reproductive function, as in real conditions organism is simultaneously exposed to great number of different chemicals, among which could be environmental pollutants, occupational factors, pharmaceuticals et cetera [10, 58]. Despite of great complicacy of such investigations, they are the key point in full understanding of real mechanisms of xenobiotics negative actions on male reproductive system. From our point of view for exact detection of all events and mechanisms, involved into development of testicular toxicity, the evaluation of cytochrome P-450 isoforms inhibitors and inducers combined action in vitro and in vivo could be very perspective. Such investigations allow working out effective methods for preventing men�s infertility under increasing pressure of adverse environmental factors, especially chemicals.
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