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© O. S. Sidorenko, G. A. Bozhok, T. P. Bondarenko UDC 612. 419. 014. 3: 611. 451
O. S. Sidorenko, G. A. Bozhok, T. P. Bondarenko
Adrenal Stem/ Progeny Cells Niches (Brief Review)
Institute for problems of cryobiology and cryomedicine of the National academy of science of Ukraine (Kharkov)
One of the areas of modern biotechnology is the use of stem cells for the treatment of several diseases. Adult tissue-specific stem cells are particularly promising in this area, because in this case there is no ethical and moral complications that are inevitable when using allogeneic cadaveric and especially abortive material. Furthermore, the use of autologous cells eliminates the need for immunosuppressive therapy directed on suppression the immune response that occurs when foreign cells and tissues are injected into the organism. The existence of a pool of tissue-specific stem / progenitor cells provides an update of the organ or tissue cell composition throughout life. These cells are also the source for the restoration of specialized differentiated cells in the case of organ damage due to injury or disease. Currently the stem cells of the liver [5, 31], pancreas [5, 29], skin [4, 30], nerve [25, 38], muscle [28, 41] etc. are described. Experimental studies aimed at developing methods for the isolation of progenitor cells from different organs, their expansion in culture conditions, as well as the potential application of them in regenerative medicine to treat a variety of pathologies are carried out in many laboratories all over the world [20].
The adrenal gland consists of two functionally distinguished endocrine tissues: the outer cortical cells, which secrete a variety of steroid hormones, and the inner medulla secreting catecholamines [1]. The adrenal gland develops from different germ layers during embryonic development: the cells of the adrenal cortex originate from the mesoderm, and the medulla cells -from the ectoderm and are neural crest derivatives [12, 21]. In the process of adrenal organogenesis spatially close of two components occurs and functional maturation of different cell types is influenced by complex paracrine interactions [7].
The adrenal gland, as the organ that develops from different germ layers, is particularly interesting in the sense that the stem / progenitor cells are found within adrenal at all levels including connective tissue capsule, the adrenal cortex and the medulla. There is close functional cooperation between differentiated cells of particular zones in the adrenal gland. This provides a harmonious work of a whole organ, which is the regulation of all types of metabolism, water-salt metabolism, participation in adaptation to various stressors.
The variety of the adrenal gland function determines the importance of this organ for the maintenance of homeostasis. Thus investigations of mechanisms that
ensure the stable operation of this organ in the living organism are of current interest.
Adrenal cortex develops as a thickening of the celomic epithelium in the cranial region of the mesonephros. Adrenogonadal primordium formed as a result of proliferation of coelomic epithelial cells and mesenchymal cells of mesonephros. Further this structure differentiates into gonadal primordium and adrenal primordium, which represents the fetal zone of the cortex at this stage. Then the mesenchymal capsule forms around the fetal cortex. As soon as the formation of the capsule is completed, definitive cortex develops between the capsule and the fetal cortex. Further the fetal cortical cells undergo apoptosis, which gradually led to its extinction during first three months of postnatal development. Human definitive adrenal cortex consists of three zones: zona glomerulosa, zona fasciculata and zona reticularis, each of which is a source of certain steroid hormones. Formation of adult adrenal zonation is completed by 20 years old and persists throughout life [16, 26]. Cells of the definitive cortex update constantly. This suggests the existence of stem/progenitor cells within adrenal and raises a lot of assumptions about their location.
As the definitive cortex is formed between the capsule and the fetal cortex it is reasonable to assume that the source of adrenocortical stem cells is either of these two structures. However recent data has allowed finding the «golden mean» and combining the two models in the form of the hypothesis assuming the existence of “niches” of adrenocortical stem / progenitor cells.
In experiments studying the adrenal cortex tissue regeneration, as well as in the labeling experiments has been shown that the most active proliferation occurs in peripheral areas of the cortex, where cells migrate centripetally and eventually die in the z. reticularis/medullary boundary [17, 39]. So the source of stem cells is either in the area located on the border of zona glomerulosa and zona fasciculata [27, 40], or mesenchymal capsule or subcapsular region [17].
A steroidogenic factor 1 (Sf-1) is important in the development of the adrenal cortex. This protein, a member of the nuclear receptor family, is an intracellular transcription factor. It regulates the transcription of key genes involved in the development of the adrenal glands, gonads, hypothalamus, and controls the production of hormones in the steroidogenic tissues [16].
Shh protein plays a key role in regulating vertebrate organogenesis. It is involved in the development of the adrenal glands, and mutations of the gene encoding the Shh synthesis, cause aplasia or abnormal development of adrenal glands [6, 19]. It was shown that cortical cells located at the periphery express Shh protein and Sf-1. At the same time, expression of Cyp11b1 or Cyp11b2 genes, which are markers of differentiated steroidogenic cells was not seen in these cells [18]. These data support the fact that the progenitor cells of the adrenal cortex are located on the periphery. However, the question of their definite localization remains controversial.
Adrenocortical cells that were transplanted into im-munodeficient mice formed adrenocortical tissue in the body of the recipient. These cells were able to respond to hormonal stimuli, but new-formed adrenocortical tissue did not have zonal structure, which were specific to the native tissue, and loss the ability to continuous cell renewal, that is, its viability decreased after serial transplantation [35]. This could be due to the lack of complex capsule I subcapsular area during transplantation. Conversely, when the entire adrenal tissue except for the capsule was removed (so-called adrenal enucleation) the complete regeneration of the cortex was possible. A similar effect of all adrenal zone cell recovery is observed during syngeneic heterotopic transplantation of adrenal capsule cells in mice [18].
However, the literature data are suggestive of the presence of undifferentiated cells directly in the adrenal cortex.
Interestingly, only 5 out of 20 randomly selected clones formed well-vascularized tissue and stabilized levels of glucocorticoids in adrenalectomized mice when xenotransplantation of clonal adrenocortical cells was performed [34]. This may be evidence for the existence of the undifferentiated cells with different proliferative potential within the adrenal cortex.
In paper [33] the authors separated zona glomeru-losa and zona fasciculata cells and transplanted them into rats with bilateral adrenalectomy. Zona glomeru-losa cells or zona fasciculata cells were transplanted to the animals of the different experimental groups, some animals were injected with a total cell suspension. Interestingly, although the viable cells of the cortex were identified in all grafts, architectonic formation specific to the adrenal cortex was observed only when zona glo-merulosa cells were present in the graft. In these animal groups the aldosterone levels were much higher than in the animals that received zona fasciculata cells only as a transplant. The authors also pointed out that in primary culture fasciculata cell number remained stationary although glomerulosa cell number increased to almost 10-fold. These data indicate that there are actively proliferating undifferentiated cells in the zona glomerulosa and confirm that the migration model in which cells from zona glomerulosa migrate centripetally and acquire zona fasciculata cells phenotype is the most correct explanation of the adrenocortical zonal structure formation.
Thus, despite the fact that the presence of progenitor cells in the peripheral region of the adrenal cortex is
no doubt, the question about more precise their localization is still not resolved. Probably stem / progenitor cells are widespread in both capsule and subcapsular region, as well as in the glomerular cortex, and vary depending on the animal species and perhaps age.
In contrast to the adrenal cortex medulla is of neuroectodermal origin and develops from the neural crest.
Neural crest is a temporary embryonic structure, which is a combination of cells released from the dorsal regions of the neural groove during its closure in the neural tube. Neural crest cells move from the neural tube, actively migrating and reaching a certain area of the developing embryo to differentiate into cells of different types, depending on the microenvironment [22]. Neural crest derivatives are found in many organs and tissues of the adult organism. Thus peripheral ganglia neurons, glial (Schwann) cells, melanocytes, some endocrine cells - adrenal chromaffin cells and thyroid C cells develop from the neural crest [24]. All of these types of cells are neural derivatives of neural crest. Chondrocytes, osteocytes, adipocytes, myofibroblasts, connective tissue cells also develop from the neural crest cells and represent its mesenchymal derivatives [24].
Chromaffin cells of the adrenal medulla is a neuroendocrine neural crest derived cells. Together with the sympathetic neurons in the spinal ganglia and small intensely fluorescent cells (which are intermediate between chromaffin cells and sympathetic neurons) they form a sympathoadrenal lineage of neural crest derivatives. It is believed that the sympathetic neurons and chromaffin cells originate from a common sympathoadrenal precursor [3, 15], which acquires appropriate properties depending on the local microenvironment.
As derivatives of common sympathoadrenal precursor, mature sympathetic neurons and adrenal chromaffin cells possess their characteristic features. Thus, chromaffin cells do not contain neurofilament and do not form processes in vivo. They express the adrenaline-synthesizing enzyme PNMT [23]. In contrast to the sympathetic neurons adrenal medulla cells retain the ability to proliferate throughout life in the adult organism [32, 37].
Given the common origin of sympathetic neurons and chromaffin cells of the adrenal glands, it has been suggested that there is cell population in fetal adrenal gland that possesses the signs of both sympathoadrenal lineage derivatives and selectively inhibits the expression of some markers, depending on the microenvironment. It has been shown that 20-40% of the rat embryo adrenal cells co-express PNMT, as well as intermediate and macromolecular neurofilament in vivo. When cultured in medium without corticosteroids the formation of neurofilament and processes was observed in 20% of bovine chromaffin cells. Moreover such cells were PNMT positive [11].
Previously it was thought that the process of adreno-medullary progenitor cells differentiation into chromaffin endocrine cell is controlled by glucocorticoids [14]. According to conventional theory, the sympathoadrenal (SA) progenitor cells develop from the neural crest cells
that are grouped near the dorsal aorta and form the primary sympathetic ganglia. Under the action of bone morphogenetic proteins (BMPs), which are secreted by the wall of the dorsal aorta, the cells acquire properties of catecholaminergic neurons. They begin to express neuronal markers and enzymes necessary for the synthesis of norepinephrine, tyrosine hydroxylase (TH) and dopamine - p - hydroxylase (DBH). Then, from the primary ganglion they migrate to their destination - the secondary sympathetic ganglia and adrenal anlage to differentiate into sympathetic neurons or chromaffin cells respectively [15]. Thus, the topography is the determining factor for neural crest cells to choose a pathway of differentiation. Cells which remain near the neural tube and the dorsal aorta, go into microenvironment rich in neural growth factor (NGF) and other specific growth factor that promotes the development of neurons in the spinal ganglia. Cells which continue the migration to the level of the adrenal gland - an area rich in glucocorticoid - differentiate into endocrine cells [14].
In the absence of glucocorticoids, some neonatal rat chromaffin cells formed processes and acquired the morphology of neuronal cells in the medium without the addition of NGF. The processes formation were observed in areas with a high content of fibroblast-like cells, which could be a source of NGF or any other growth factors in this case [9, 10].
In vitro cell culture studies support the hypothesis that considers glucocorticoids as a major factor in the differentiation of sympathoadrenal precursor into chromaffin cells. It was shown that glucocorticoids are necessary for the manifestation of the characteristic properties of adrenal chromaffin cells in culture. In a medium without glucocorticoids the activity of PNMT (the enzyme that convert noradrenaline molecules into adrenaline molecules and is one of the markers of chromaffin cells) decreased completely. As a result the synthesis and accumulation of adrenaline decreased, although the amount of catecholamines in general remained at the level of control values. Moreover, cell survival decreased dramatically in the absence of glucocorticoids during long-term culturing [9]. The decrease in viability was characteristic of cells derived from both neonatal and adult animals, although the latter was less expressed.
Recently, however, the paramount importance of glucocorticoids for the formation of chromaffin cell phenotype is debatable. This is associated with breeding lines of mice carrying a mutation of the gene responsible for the synthesis of glucocorticoid receptor [13]. The adrenal medulla in these mice developed normally, despite the absence of glucocorticoid receptors.
It should be noted that the differentiation of adre-nomedullary cell precursors into chromaffin cells is not final and is reversible: the transcription of neurospecific genes that were expressed during ontogeny can begin under the influence of certain stimuli [14]. This reflects the chromaffin cells ‘’plasticity’’. One of the most studied factors that stimulate expression of neuronal phenotype in the mature adrenomedullary cells is NGF
Cells of the adrenal medulla are beginning to form neurofilaments, processes with synaptic contacts and generate an action potential when NGF was added to the culture medium [8, 9]. Many scientists study the NGF-induced phenotypic changes of adrenal medulla cells. However it remains debatable the initial degree of differentiation of cells undergoing NGF- induced phenotype changes. Whether they are multipotent progenitor cells or mature chromaffin cells? In the first case it is the final differentiation into neuronal cells under the influence of NGF and in the second case it should be referred as transdifferentiation. In addition, these cells may belong to a small population of medullary neurons that are originally present in the medulla and grow in culture supplemented with NGF. The last assumption is unlikely because adrenal medulla cells has been cultured for a week without NGF [9], while the sympathetic ganglia neurons die within 24-48 hours without NGF [36].
The authors [9] have shown that the adrenal medulla cells that changed their phenotype into neurons under the influence of NGF, was originally a fully differentiated chromaffin cells. This was evidenced by morphological analysis, an intense catecholamine fluorescence (characteristic of adrenal chromaffin cells, but not neurons [9, 10]), and the high level of PNMT activity. When cultured in glucocorticoid-free medium enriched with NGF, catecholamine fluorescence disappeared completely, although half of the cells, which were seeded, remained in a viable state. Moreover the PNMT activity decreased completely.
However, in other studies the authors observed the differentiation of adrenal medulla progenitor cells into the neurons under the action of NGF. Adrenomedullary cells have been cultured for two weeks in conditions that prevented their attachment to the surface. In addition, the steroid-free culture medium was used. In these conditions, the majority of chromaffin cells died, and the few surviving cells formed spherical floating colonies - chromospheres. Analysis of the cells that constitute the chromosphere showed that expression of PNMT decreased 100-fold, and the expression of nes-tin - a marker of neural stem cells - increased by 6-fold compared to cells before culturing.
The differentiation of these progenitor cells into the nervous and endocrine direction induced by NGF and dexamethasone, respectively. The cells did not express PNMT before stimulation of differentiation.
The NGF administration stimulated formation of processes containing III p-tubulin. 24% of the cells generate action potentials followed by hyperpolarization after differentiation. Since most of the differentiated chromaffin cells died in these contrary conditions, the culture enriched with progenitor cells was obtained. Most likely, neuron cells in this case were the result of progenitor cells differentiation rather than chromaffin cells transdifferentiation.
It was shown that there is a population of cells in the newborn piglets adrenal glands that forms multicellular structures and subsequent differentiation of cells into neurons is observed. It is interesting that the conditions needed for the formation of such structures and cell
differentiation differ from those described previously for Thus, all parts of the adrenal gland have its own pool the chromosphere: multicellular structures not formed of stem I progenitor cells, which ensure the constancy of in the suspension but on the monolayer only, and neu- adrenal cellular composition during the whole life of the
ronal differentiation does not require the introduction of organism. Probably, the potentia|ities of adrena| stem
. А, , A .. А| A , А| cells in regenerative medicine are very large. Therefore
any external growth factors. However the nature of the xl . ,
the number of questions regarding the mechanisms
ce||s that form mu|tice||u|ar structures, their |oca|ization and factors of stem cells differentiation, methods for
within the adrenal gland, as well as the degree of their their isolation, culturing and expansion in vitro, which is
lineage commitment and, therefore, the possibility to especially important in the case of their application for
differentiation remain a challenging open problem [2]. autologous transplantation, remain actual.
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УДК 612. 419. 014. 3: 611. 451
НИШИ СТВОЛОВЫХ/ПРОГЕНИТОРНЫХ КЛЕТОК НАДПОЧЕЧНИКОВ (краткий обзор литературы) Сидоренко О. С., Божок Г. А., Бондаренко Т. П.
Резюме. Гормоны надпочечников контролируют множество биохимических и физиологических процессов в организме человека, поэтому актуальной задачей современной медицины является исследование механизмов, обеспечивающих постоянство клеточного состава и стабильную работу самого органа. В кратком обзоре литературы представлены основные предположения относительно областей локализации стволовых прогенитрных клеток надпочечников и возможности их использования в регенеративной медицине.
Ключевые слова: стволовые клетки надпочечников, капсула, кора надпочечников, медулла, регенеративная медицина.
УДК 612. 419. 014. 3: 611. 451
НИШІ СТОВБУРОВИХ/ПРОГЕНІТОРНИХ КЛІТИН НАДНИРНИКІВ (стислий літературний огляд) Сидоренко О. С., Божок Г. А., Бондаренко Т. П.
Резюме. Гормони надниркових залоз контролюють велику кількість біохімічних та фізіологічних процесів в організмі людини, тому актуальним питанням сучасної медицини є дослідження механізмів, що забезпечують сталість клітинного складу та стабільну роботу самого органа. В стислому огляді літератури представлено основні припущення щодо сайтів локалізації стовбурових!прогеніторних клітин наднирників та перспективи їх використання в регенеративній медицині.
Ключові слова: стовбурові клітини наднирників, капсула, кора наднирників, медула, регенеративна медицина.
UDC 612. 419. 014. 3: 611. 451
Adrenal Stem/ Progeny Cells Niches (Brief Review)
Sidorenko O. S., Bozhok G. A., Bondarenko T. P.
Summary. Adrenal hormones control a lot of biochemical and physiological processes in the human body, so the actual problem of modern medicine is the study of mechanisms that ensure the constancy of cellular composition and stable functioning of the adrenal gland. In a brief review the main assumptions about adrenal stemIprogeny cells localization and prospects of their application in regenerative medicine are presented.
Key words: adrenal stem cells, capsule, adrenal cortex, medulla, regenerative medicine.
Стаття надійшла 28. 02. 2013 р. Рецензент - проф. Розанов Л. Ф.
