Nature, significance, and cytological consequences of cytomixis

Cytomixis is a widespread natural process of intercellular interaction, which is common for vegetative and generative tissues in both normal and pathological conditions. The origin, significance, and genetic control of cytomixis still remain partially unclear. The widespread point of view regarding the pathological nature of cytomixis is based on its singularity for plants with genetic instability and impaired homeostasis. Meiotic genes responsible for the segregation and organization of chromosomes, the activity of which is modified by environmental factors through the signal transduction system, seem to be involved in the genetic control of cytomixis. It is assumed that, via cytomoxis, on the one hand, an information contact can be reached as well as meiosis and gametogenesis are synchronized and, on the other hand, the genetic variety and heterozygosis of microsporocytes increase. The activity of cytomoxis varies within a wide interval. Mutagenesis, hybridization, induction, and polyploidy have the greatest influence on this activity. In this context, cytomixis may have a function of cell selection, which is activated by exceeding the threshold level of microsporocytes damage (or genetic disbalance).     

New insights into cytomixis: specific cellular features and prevalence in higher plants

The phenomenon of intercellular migration of nuclei in plant tissues (cytomixis) was discovered over a century ago, which has been followed by numerous attempts to clarify the essence of this process as well as to determine its causes and consequences. Most attention of researchers has been paid to cytomixis in microsporogenesis, since the transfer of part of genetic material between microsporocytes may influence the ploidy level of the produced pollen and, presumably, have an evolutionary significance. This review compiles the data on cytological pattern of cytomixis and proposes a scheme as to how cytomictic channels are formed and function in angiosperms. The prevalence of cytomixis in different plant taxa is analyzed using the published data. The causes, mechanisms, and consequences of the nuclear migration between cells in plant tissues are discussed.     

Impact of cytomixis on meiosis, pollen viability and pollen size in wild populations of Himalayan poppy (Meconopsis aculeata Royle)

We report the occurrence of cytomixis in wild populations of Himalayan poppy (Meconopsis aculeata Royle), which is considered to be an important and threatened medicinal plant growing in the high hills of the Himalayas. The impact of cytomixis on meiotic behaviour, reduced pollen viability and heterogeneous-sized pollen grains was also studied. Cytological studies in the seven wild populations from the high hills of Himachal Pradesh revealed that all the Himalayan populations exist uniformly at the tetraploid level (2n=56) on x=14. The phenomenon of chromatin transfer among the proximate pollen mother cells (PMCs) in six populations caused various meiotic abnormalities. Chromatin transfer also resulted in the formation of coenocytes, aneuploid, polyploid and anucleated PMCs. Among individuals that showed chromatin transfer, chromosome stickiness and interbivalent connections were frequently observed in some PMCs. The phenomenon of cytomixis in the species seems to be directly under genetic control; it affects the meiotic course considerably and results in reduced pollen viability.     

An ultrastructural study of cytomixis in tobacco pollen mother cells

Intercellular chromatin migration (cytomixis) in the pollen mother cells of two tobacco (Nicotiana tabacum L.) lines was analyzed by electron microscopy during the first meiotic prophase. The maximal manifestation of cytomixis was observed in the pachytene. As a rule, several cells connected with one another by cytomictic channels wherein the nuclei migrated were observable at this stage. In the majority of cases, nuclei passed from cell to cell concurrently through several closely located cytomictic channels. Chromatin migrated between cells within the nuclear envelope, and its disintegration was unobservable. The nucleus, after passing through cytomictic channels into another cell, can be divided into individual micronuclei or, in the case of a direct contact with another nucleus, can form a nuclear bridge. It has been demonstrated that the chromatin structure after intracellular migration visually matches the chromatin structure before it passed through the cytomictic channel. No signs of pyknosis were observable in the chromatin of the micronuclei formed after cytomixis, and the synaptonemal complex was distinctly seen. The dynamics of changes in the nucleoli during cytomixis was for the first time monitored on an ultrastructural level. Possible mechanisms determining cytomixis are discussed and the significance of this process in plant development is considered.     

Methomyl,imbraclaobrid and clethodim induced cytomixis and syncytes behaviors in PMCs of Pisum sativum L: Causes and outcomes

Cytomixis is a common phenomenon observed in meiotic cells such as anther which is influenced by various factors. Use of pesticides is a common practice in agriculture. However, it is not known whether pesticides can induce cytomixis in plant cells and induce genetic variation. To understand this, the present study was planned to assess the cytomixis and syncytes behaviors in PMCs of Pisum sativum L. Seeds of P. sativum (Family: Fabaceae) were treated with different concentrations of commonly used pesticides methomyl (ME), imbraclaobrid (IM) and clethodim (CL). Seeds were treated with various concentrations (0.1, 0.2, 0.3, 0.4 and 0.5% of ME, IM and CL prepared in water) for 1 and 3 h. Effect of pesticides on pollen fertility, frequency of cytomixis, and kind of cytomixis cells was assessed. In the cytomixis cells, the cytomictic channel (CC) and direct fusion (DF), and various stages of meiosis (PI, MI, AI and TI) with cytomixis cells were observed. In addition, frequency of syncytes cell and their various stages of meiosis I (PI, MI, AI and TI) in pollen mother cells (PMCs) was assessed. During the microsporogenesis in P. sativum, the occurrence of cytomixis and syncytes at various stages of meiosis I were seen. The formation of cytoplasmic channels and direct fusing of pollen mother cells (PMCs) were both seen to cause cytomixis, with the former being more common than the latter. The percentage of PMCs with cytomixis and syncytes cells increased with increase in the concentration of pesticides. The result of the present investigation indicates that commonly used pesticides ME, IM, and CL have a significant effect on pollen fertility, frequency of cytomixis, and kind of cytomixis cells, the cytomictic channel (CC) and direct fusion (DF), in addition, frequency of syncytes cell and their various stages of meiosis I (PI, MI, AI and TI) in pollen mother cells (PMCs) on P. sativum.     

Cytomixis and its possible evolutionary role in a Kuwaiti population of Diplotaxis harra (Brassicaceae)

During a cytotaxonomical study of a Kuwaiti diploid (2 n  = 26) population of Diplotaxis harra (Forssk.) Boiss (Brassicaceae), cytomixis and aneuploidy were found in 1.5% and 7.8% of the pollen mother cells (PMCs), respectively. Cytomictic cells revealed wide variability as far as the number of involved PMCs and their stage of division were concerned. The cytoplasmic channels, which were of different sizes, contained migrating chromosomes in a few cases. In some PMCs, the location of some of the chromosomes was clearly indicative of an imminent migration from a donor to its attached recipient cell. At metaphase I and anaphase I, PMCs showed variable degree of chromosomal transfer. In addition to a genetic control of the phenomenon, stress factors such as high temperature or drought in certain periods of the growing season may have contributed to the incidence of cytomixis in the material. A possible relationship between cytomixis and detected aneuploid PMCs, as well as a reconsideration of an evolutionary role of cytomixis is discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 169–175.     

Cytomixis in plants: facts and doubts

The migration of nuclei between plant cells (cytomixis) is a mysterious cellular phenomenon frequently observable in the male meiosis of higher plants. Cytomixis attracts attention because of unknown cellular mechanisms underlying migration of nuclei and its potential evolutionary significance, since the genetic material is transferred between the cells that form pollen. Although cytomixis was discovered over a century ago, the advance in our understanding of this process has been rather insignificant because of methodological difficulties. The data that allowed for a new insight into this phenomenon were obtained by examining the migrating nuclei with electron and confocal laser microscopy, immunostaining, and fluorescence in situ hybridization. As has been shown, the chromatin migrating between cells is surrounded by an undamaged nuclear membrane. Such chromatin does not undergo heterochromatization and contains normal euchromatin markers. The condensation degree of the migrating chromatin corresponds to the current meiotic stage, and normal structures of synaptonemal complex are present in the migrating part of the nucleus. The cells involved in cytomixis lack any detectable morphological and molecular markers of programmed cell death. It has been shown that individual chromosomes and genomes (in the case of allopolyploids) have no predisposition to the migration between cells, i.e., parts of the nucleus are involved in cytomixis in a random manner. However, the fate of migrating chromatin after it has entered the recipient cell is still vague. A huge amount of indirect data suggests that migrating chromatin is incorporated into the nucleus of the recipient cell; nonetheless, the corresponding direct evidences are still absent. No specific markers of cytomictic chromatin have been yet discovered. Thus, the causes and consequences of cytomixis are still disputable. This review briefs the recent data on the relevant issues, describes the classical and modern methodological approaches to analysis of the intercellular migration of nuclei, and discusses the problems in cytomixis research and its prospects.     

Cytomixis and its role in the regulation of plant fertility

UV and gamma irradiation of barley seedlings induces an increase in the number of various pathologies in the male reproductive system of plants. The majority of cytological abnormalities are rather nonspecific. The main type of the observed pathologies of microsporogenesis is cytomixis, whose activation correlates with a callose hypersecretion in microsporocyte walls. A negative correlation between cytomixis and the sterility of microspores (in the case of gamma irradiation) or the sterility of mature pollen grains (in the case of UV-B irradiation) is revealed. It is supposed that cytomixis represents a kind of a premeiotic cell selection in plants characterized by an intraorganismic genetic heterogeneity (mosaics). The novelty of the idea is that the cytopathology that accompanies cytomixis is considered as a mechanism of the induced death of genetically imbalanced or nonrepairable cells, which is intended to keep the fertility of a male reproductive system. The activation of this mechanism has a threshold character.     

A rise of ploidy level influences the rate of cytomixis in tobacco male meiosis

The effect of plant ploidy level on the rate of cytomixis in microsporogenesis has been analyzed with the help of a unique model, the collection of tobacco plants of different ploidies (2n?=?2x?=?24, 4x?=?48, 6x?=?72, and 8x?=?96). As has been shown, the rate of cytomixis proportionally increases in 6x and 8x cytotypes, being rather similar in 2x and 4x plants. The rate of cytomixis is highly variable, differing even in the genetically identical plants grown under the same conditions. The cytological pattern of cytomixis in the microsporogenesis of control 4x plants has been compared with the corresponding patterns of 2x, 6x, and 8x plants. Involvement of cytomixis in production of unreduced gametes and stabilization of the newly formed hybrid and polyploidy genomes is discussed.     

百合花粉母细胞间细胞融合期间腺苷三磷酸酶活性的细胞化学定位及其与染色质胞间转移的关系

用标准的磷酸铅沉淀的细胞化学方法,对百合花粉母细胞间染色质穿壁运动期间及其前后三个时期中的腺苷三磷酸酶(ATP 酶)活性进行了超微结构的定位。结果表明:(1)在穿壁前,ATP 酶活性主要定位于质膜、胞间连丝及细胞间隙;在内质网、高尔基体、质体和某些局部的基质(groundplasm)中,也表现有 ATP 酶活性反应的产物;但在染色质和核仁中,一般都没有这种反应。(2)在穿壁时,染色质从一个细胞穿壁转移到另一个相邻细胞,同时看到染色质和核仁内出现密集的 ATP 酶活性反应产物;在内质网和高尔基体的腔内以及质体的片层上也产生明显的 ATP 酶活性反应;而在质膜、胞间连丝及细胞间隙内 ATP 酶活性明显降低,甚至看不到明显的活性反应。(3)在穿壁后,质膜及细胞间隙中又产生明显的 ATP 酶活性反应产物,但核内染色质上的 ATP 酶活性则显著降低,而核仁内则仍有较高的活性。同前二个时期一样,内质网、高尔基体和质体上的 ATP 酶仍表现明显的活性反应。最后讨论了三个不同发育时期 ATP 酶活性及其分布部位的改变与染色质胞间转移的关系。     

Efficiency of the induction of cytomixis in the microsporogenesis of dicotyledonous (<Emphasis Type="Italic">N. tabacum</Emphasis> L.) and monocotyledonous (<Emphasis Type="Italic">H. distichum</Emphasis> L.) plants by thermal stress

The efficiencies of the induction of cytomixis in microsporogenesis by thermal stress are compared in tobacco (N. tabacum L.) and barley (H. distichum L.) It has been shown that different thermal treatment schedules (budding tobacco plants at 50°C and air-dried barley grains at 48°C) produce similar results in the species: the frequency of cytomixis increases, and its maximum shifts to later stages of meiosis. However, the species show differences in response. The cytomixis frequency increase in tobacco is more pronounced, and its maximum shifts from the zygotene–pachytene stages of meiotic prophase I to prometaphase–metaphase I. Later in the meiosis, aberrations in chromosome structure and meiotic apparatus formation typical of cytomixis are noted, as well as cytomixis activation in tapetum cells. Thermal stress disturbs the integration of callose-bearing vesicles into the callose wall. Cold treatment at 7°C does not affect cytomixis frequency in tobacco microsporogenesis. Incubation of barley seeds at 48°C activates cytomixis in comparison to the control, shifts its maximum from the premeiotic interphase to zygotene, and changes the habit of cytomictic interactions from pairwise contacts to the formation of multicellular clusters. Thermal treatment induces cytomictic interactions within the tapetum and between microsporocytes and the tapetum. However, later meiotic phases show no adverse consequences of active cytomixis in barley. It is conjectured that heat stress affects callose metabolism and integration into the forming callose wall, thereby causing incomplete closure of cytomictic channels and favoring intercellular chromosome migration at advanced meiotic stages.     

Cytochemical Localization of Adenosine Triphosphataes Activity During Cytomixis in Pollen Mother Cells of David Lily and Its Relation to the Intercellular Migrating Chromatin Substance

Standard lead precipitation procedures have been used to examine the localization of ATPase activity during cytomixis in pollen mother cells of Lilium davidii var. willmottiae (Wilson) Roffill. Before cytomixis, cells at this stage of development show ATPase activity on plasma membrane, in the endoplasmic reticulum, dictyosomes, plastids, plasmodesmata, and in part of the groundplasm; however, there is no ATPase activity on the chromatin and nucleolus. During cytomixis, the chromatin substance begin to transfer from one cell to an adjacent cell, reaction product indicating ATPase activity is observed associated with the chromatin and nucleolus. ATPase activity is also found with the cistenae of both endoplasmic reticulum and dictyosomes, and some plastids. There is no deposition of ATPase reaction product associated with the plasm membrane and intercellular spaces. After cytomixis, the chromatin is little or no deposition of enzyme reaction product. ATPase activity, however, is consistenlly found within the intercellular space and on the plasm membrane, and also occur in the endoplasmic reticulum, dictyosome and plastid. The presence or absence of ATPase activity in the cell structure of pollen mother cells before, during or after eytomixis is discussed in relation to the active uptake or export of water for short-distance transport. It is also suggested that the intensive ATPase activity in the nucleus during cytomixis of pollen mother cells is evidence for a transport system involved in the active movement of the intercellular migrating ebromatin substance.     

Cytomixis in pollen mother cells of Medicago sativa L

Cytomixis (i.e., chromatin migration between meiocytes) has been detected in many plant species, but not in Medicago sativa spp. In the present study we report the identification of a few cytomictic alfalfa plants. Those plants, the "mother plants," were selfed and crossed with a normal control plant. Microsporogenesis analysis was performed on the mother plants, on the S(1) and F(1) plants, and on controls. The S(1) and F(1) plants, like the mother plants, were found to be cytomictic. Single or multiple chromatin bridges between two or more meiocytes were observed almost exclusively in prophase I. Some completely empty meiocytes were also observed. In addition to cytomixis, other meiotic abnormalities were found. Control plants showed an almost regular meiosis. The highest values of cytomixis were observed in the mother plants, and the lowest in their F(1) progenies. Variability of cytomixis in the F(1) plants is probably due to a heterozygotic condition of the parents for this trait. No significant correlation was found between cytomixis and pollen viability, even if the cytomictic plants showed low values of pollen viability.     

First detection of cytomixis and its consequences in <Emphasis Type="Italic">Thalictrum cultratum</Emphasis> Wall. (Ranunculaceae)

In current paper, we studied the meiotic chromosome number and details of cytomixis recorded for the first time in Thalictrum cultratum Wall. from the high altitude areas of northwest Himalaya (India). Cytomixis, the phenomenon of inter PMC (pollen mother cell) migration of chromatin material has been reported in many angiosperms and other groups of plants, but there is no published report in Thalictrum cultratum. All the presently studied accessions of the species existed at diploid level (x = 7). The present chromosome count of n = 14 has been reported for the first time from the study area. The cytological stability of any plant is an important consideration in view of its existence in nature. Meiotic disturbances can impose threat to the reproductive potential of plant. Current study reveals the occurrence of inter PMC migration of chromatin material. In present investigation, the phenomenon of cytomixis can be observed between 2 to 9 PMCs. Chromatin migration occurs through narrow and broad cytoplasmic channels or occasionally fused together during metaphase-I and anaphase-I. Cytomixis resulted in the formation of hypo- (n = 2, 2 + 1 small fragment, n = 3 + 3 small fragments, 4, 5 + extra mass of chromatin, 13) and hyperploid (n = 7 + 2 small fragments, 8 + 2 small fragments, 14, 14 + 2 small fragments, 15) PMCs, variable sized pollen grains and pollen sterility (20%). The current study reveals that cytomixis is responsible for formation of PMCs with variable chromosomes (hypoand hyperploid) and pollen grains of heterogeneous sizes and pollen sterility in T. cultratum. This is the first report of cytomixis in the species.     

The role of microtubular cytoskeleton and callose walls in the cytomixis process in tobacco (Nicotiana tabacum L.) pollen mother cells

We have studied the microtubule cytoskeleton structure and callose walls deposition in the course of meiosis at the cytomictic and normal tobacco (N. tabacum L.) PMCs. It was ascertained that microtubule cytoskeleton did not play an evident part in the process of cytomixis. Increased cytomixis frequency probably is determined by irregular callose walls deposition. The possible reasons of nuclear material passage between tobacco PMCs at the cellular level are discussed.     

Effect of N-stearoylethanolamine on the level of stable NO metabolites in different pathological conditions which are accompanied by oxidative stress

The effects of N-stearoylethanolamine (NSE) at the level of stable NO metabolites--NO2- NO3- under different pathological conditions which were accompanied by oxidative stress and NO disbalance were studied. It was found that NSE promoted the increasing of NO metabolites level on the animal models with deficit of NO compared to control and evoked decreasing content of NO2- and NO3- under pathological conditions with NO overproduction. It is suppoused, that NSE effects could be determined, by its ability to modulate the the activity of NO-synthase izoformes.     

Induced cytomictic variations and syncyte formation during microsporogenesis in <Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> L.

The intercellular translocation of chromatin material along with other cytoplasmic contents among the proximate meiocytes lying in close contact with each other commonly referred as cytomixis was reported during microsporogenesis in Phaseolus vulgaris L., a member of the family Fabaceae. The phenomenon of cytomixis was observed at three administered doses of gamma rays viz. 100, 200, and 300 Gy respectively in the diploid plants of Phaseolus vulgaris L. The gamma rays irradiated plants showed the characteristic feature of inter-meiocyte chromatin/chromosomes transmigration through various means such as channel formation, beak formation or by direct adhesion between the PMC’s (Pollen mother cells). The present study also reports the first instance of syncyte formation induced via cytomictic transmigration in Phaseolus vulgaris L. Though the frequency of syncyte formation was rather low yet these could play a significant role in plant evolution. It is speculated that syncyte enhances the ploidy level of plants by forming 2n gametes and may lead to the production of polyploid plants. The phenomenon of cytomixis shows a gradual inclination along with the increasing treatment doses of gamma rays. The preponderance of cytomixis was more frequent during meiosis I as compared to meiosis II. An interesting feature noticed during the present study was the channel formation among the microspores and fusion among the tetrads due to cell wall dissolution. The impact of this phenomenon is also visible on the development of post-meiotic products. The formation of heterosized pollen grains; a deviation from the normal pollen grains has also been reported. The production of gametes with unbalanced chromosomes is of utmost importance and should be given more attention in future studies as they possess the capability of inducing variations at the genomic level and can be further utilized in the improvement of germplasm.     

The indication of genetic and environmental components in behaviour

The problem of genetic influences on behaviour was examined under the aspect of inter-specific differences, which are predomintantly genetic and are the basis of sociobiology. It was also examined under the aspect of intra-specific differences, notably the normal differences of behaviour which, like the sex differences, have a genetic basis. Further evidence for possible genetic influences on behaviour was given by considering limiting behaviours to which a limiting phenotype, usually pathological, corresponds. A comparison between genetic and non-genetic phenomena at the population level, shows a marked difference in the temporal variation of the two types of phenomena: those wich are under the control of gene frequencies tend to remain constant whereas those which are not under genetic control vary widely from one generation to the next, or from one year to the next. In Italy, in the period 1887–1980, it was observed that frequency of homicidal cases was extremely variable in time; whereas suicide in the same period was much less variable, and seemed correlated with genetic entities. It was reasoned that, although the study of human differences, including genetic ones, is amply desirable, the variations of extreme anomic behaviours are not demonstrably under genetic control; if a genetic component exists for such behaviours it must be quantitatively very small.