Germ-Soma Differentiation

Last updated

Germ-Soma Differentiation is the process by which organisms develop distinct germline and somatic cells. The development of cell differentiation has been one of the critical aspects of the evolution of multicellularity and sexual reproduction in organisms. Multicellularity has evolved upwards of 25 times, [1] and due to this there is great possibility that multiple factors have shaped the differentiation of cells. There are three general types of cells: germ cells, somatic cells, and stem cells. Germ cells lead to the production of gametes, while somatic cells perform all other functions within the body. Within the broad category of somatic cells, there is further specialization as cells become specified to certain tissues and functions. In addition, stem cell are undifferentiated cells which can develop into a specialized cell and are the earliest type of cell in a cell lineage. [2] Due to the differentiation in function, somatic cells are found only in multicellular organisms, as in unicellular ones the purposes of somatic and germ cells are consolidated in one cell.

Contents

All organisms with germ-soma differentiation are eukaryotic, and represent an added level of specialization to multicellular organisms. Pure germ-soma differentiation has developed in a select number of eukaryotes (called Weismannists), included in this category are vertebrates and arthropods- however land plants, green algae, red algae, brown algae, and fungi have partial differentiation. [3] While a significant portion of organisms with germ-soma differentiation are asexual, this distinction has been imperative in the development of sexual reproduction; the specialization of certain cells into germ cells is fundamental for meiosis and recombination.

Weismann barrier

The strict division between somatic and germ cells is called the Weismann barrier, in which genetic information passed onto offspring is found only in germ cells. This occurs only in select organisms, however some without a Weismann barrier do present germ-soma differentiation. These organisms include land plants, many algaes, invertebrates, and fungi whose germ cells are derived from prior somatic cells as opposed to stem cells. The Weismann barrier is essential to the concept of an immortal germline, which passes down genetic information through designated germ cells.

Organisms with germ-soma differentiation but no Weismann barrier often reproduce through somatic embryogenesis.

Benefits and Detriments of Differentiation

There is no single widely accepted theory on the origins of somatic-germline differentiation, however of those that do exist many are based on the evolutionary advantage of differentiated multicellularity which has allowed it to survive. These theories include the development of colonial organization structures in which the division of labor between cells allowed for improvements in fitness.

The division of labor within multicellular organisms can offer significant advantages over unicellular counterparts. Division can allow organisms to become larger, or interact with the environments (and thus fill different niches) that increase fitness. In addition to internal benefits, there is evidence that these also improve defenses against predation. [4] On the other hand, multicellularity comes with increased energy use devoted to maintaining homeostasis instead of to reproduction.

Dirty Work Hypothesis

One major theory as to the proliferation of organisms with cell differentiation is the dirty work hypothesis. This hypothesis posits that when an organism has differentiated cells, somatic cells are able to devote energy solely to maintaining homeostasis instead of reproduction while germ cells do the opposite. One reason proposed for the relative success of the "dirty work" system of organization is that it helps manage the detrimental effects of metabolic activity, and allow for more efficient energy distribution throughout an organism. [5] The other major reason proposed is that it prevents metabolic activity within the cell from damaging genetic material. Said activity in mitochondria and chloroplasts creates mutagenic byproducts, so in organisms with differentiation where germ cells do not engage in metabolic activity the germline is not impacted. [5] [6]

Uncertainty

Due to the nature of research around the origin of life and multicellularity, it has been difficult to obtain a case study that is optimal for observing somatic-germline differentiation. One case that has been extensively studied is that of organisms in the Volvocacaeae family. Within volvocavea, there is much diversity in organizational structure, with some organisms being unicellular, colonial, or (arguably) multicellular. [7] Within volvocine algae three genes have been identified as crucial to the development of soma cells which regulate coding for asymmetric division of cells, preventing reproductive development of soma cells, and preventing the development of somatic characteristics in germ cells (such as those meant for mobility or metabolic activity [8] ).

Related Research Articles

<i>Volvox</i> Genus of algae

Volvox is a polyphyletic genus of chlorophyte green algae in the family Volvocaceae. Volvox species form spherical colonies of up to 50,000 cells, and for this reason they are sometimes called globe algae. They live in a variety of freshwater habitats, and were first reported by Antonie van Leeuwenhoek in 1700. Volvox diverged from unicellular ancestors approximately 200 million years ago.

<span class="mw-page-title-main">August Weismann</span> German evolutionary biologist (1834–1914)

August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after Charles Darwin. Weismann became the Director of the Zoological Institute and the first Professor of Zoology at Freiburg.

<span class="mw-page-title-main">Cellular differentiation</span> Developmental biology

Cellular differentiation is the process in which a stem cell changes from one type to a differentiated one. Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a multicellular organism as it changes from a simple zygote to a complex system of tissues and cell types. Differentiation continues in adulthood as adult stem cells divide and create fully differentiated daughter cells during tissue repair and during normal cell turnover. Some differentiation occurs in response to antigen exposure. Differentiation dramatically changes a cell's size, shape, membrane potential, metabolic activity, and responsiveness to signals. These changes are largely due to highly controlled modifications in gene expression and are the study of epigenetics. With a few exceptions, cellular differentiation almost never involves a change in the DNA sequence itself. However, metabolic composition does get altered quite dramatically where stem cells are characterized by abundant metabolites with highly unsaturated structures whose levels decrease upon differentiation. Thus, different cells can have very different physical characteristics despite having the same genome.

In cellular biology, a somatic cell, or vegetal cell, is any biological cell forming the body of a multicellular organism other than a gamete, germ cell, gametocyte or undifferentiated stem cell. Somatic cells compose the body of an organism and divide through mitosis.

In cellular biology, the term somatic is derived from the French somatique which comes from Ancient Greek σωματικός, and σῶμα is often used to refer to the cells of the body, in contrast to the reproductive (germline) cells, which usually give rise to the egg or sperm. These somatic cells are diploid, containing two copies of each chromosome, whereas germ cells are haploid, as they only contain one copy of each chromosome. Although under normal circumstances all somatic cells in an organism contain identical DNA, they develop a variety of tissue-specific characteristics. This process is called differentiation, through epigenetic and regulatory alterations. The grouping of similar cells and tissues creates the foundation for organs.

<span class="mw-page-title-main">Multicellular organism</span> Organism that consists of more than one cell

A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- and partially multicellular, like slime molds and social amoebae such as the genus Dictyostelium.

<span class="mw-page-title-main">Biological life cycle</span> Series of stages of an organism

In biology, a biological life cycle is a series of stages of the life of an organism, that begins as a zygote, often in an egg, and concludes as an adult that reproduces, producing an offspring in the form of a new zygote which then itself goes through the same series of stages, the process repeating in a cyclic fashion.

<span class="mw-page-title-main">Germ cell</span> Gamete-producing cell

A germ cell is any cell that gives rise to the gametes of an organism that reproduces sexually. In many animals, the germ cells originate in the primitive streak and migrate via the gut of an embryo to the developing gonads. There, they undergo meiosis, followed by cellular differentiation into mature gametes, either eggs or sperm. Unlike animals, plants do not have germ cells designated in early development. Instead, germ cells can arise from somatic cells in the adult, such as the floral meristem of flowering plants.

<span class="mw-page-title-main">Germline</span> Population of a multicellular organisms cells that pass on their genetic material to the progeny

In biology and genetics, the germline is the population of a multicellular organism's cells that develop into germ cells. In other words, they are the cells that form gametes, which can come together to form a zygote. They differentiate in the gonads from primordial germ cells into gametogonia, which develop into gametocytes, which develop into the final gametes. This process is known as gametogenesis.

<span class="mw-page-title-main">Weismann barrier</span> Distinction between germ cell lineages producing gametes and somatic cells

The Weismann barrier, proposed by August Weismann, is the strict distinction between the "immortal" germ cell lineages producing gametes and "disposable" somatic cells in animals, in contrast to Charles Darwin's proposed pangenesis mechanism for inheritance. In more precise terminology, hereditary information moves only from germline cells to somatic cells. This does not refer to the central dogma of molecular biology, which states that no sequential information can travel from protein to DNA or RNA, but both hypotheses relate to a gene-centric view of life.

<span class="mw-page-title-main">Germ plasm</span> Biological concept

Germ plasm is a biological concept developed in the 19th century by the German biologist August Weismann. It states that heritable information is transmitted only by germ cells in the gonads, not by somatic cells. The related idea that information cannot pass from somatic cells to the germ line, contrary to Lamarckism, is called the Weismann barrier. To some extent this theory anticipated the development of modern genetics.

Enquiry into the evolution of ageing, or aging, aims to explain why a detrimental process such as ageing would evolve, and why there is so much variability in the lifespans of organisms. The classical theories of evolution suggest that environmental factors, such as predation, accidents, disease, and/or starvation, ensure that most organisms living in natural settings will not live until old age, and so there will be very little pressure to conserve genetic changes that increase longevity. Natural selection will instead strongly favor genes which ensure early maturation and rapid reproduction, and the selection for genetic traits which promote molecular and cellular self-maintenance will decline with age for most organisms.

<i>Gonium</i> Genus of algae

Gonium is a genus of colonial algae, a member of the order Chlamydomonadales. Typical colonies have 4 to 16 cells, all the same size, arranged in a flat plate, with no anterior-posterior differentiation. In a colony of 16 cells, four are in the center, and the other 12 are on the four sides, three each. A description by G.M. Smith :

Gonium Mueller 1773: Colonies of 4-8-16 cells arranged in a flat quadrangular plate and embedded in a common gelatinous matrix or connected by broad gelatinous strands. Cells ovoid to pyriform, with a single cup-shaped chloroplast containing one pyrenoid. Each cell with two cilia of equal length, contractile vacuoles at the base of the cilia, and an eyespot. Four- and eight-celled colonies with the cilia on the same side ; sixteen-celled colonies with the four central cells having their cilia on the same side and the twelve marginal cells with radially arranged cilia.

Asexual reproduction by simultaneous division of all cells in the colony to form autocolonies, or by a formation of 2-4 zoospores in each cell.

Sexual reproduction isogamous, by a fusion of biciliatezoogametes.

<span class="mw-page-title-main">Udoteaceae</span> Family of algae

Udoteaceae is a family of green algae, in the order Bryopsidales.

<span class="mw-page-title-main">Outline of cell biology</span> Overview of and topical guide to cell biology

The following outline is provided as an overview of and topical guide to cell biology:

<span class="mw-page-title-main">Deep homology</span> Control of growth and differentiation by deeply conserved genetic mechanisms

In evolutionary developmental biology, the concept of deep homology is used to describe cases where growth and differentiation processes are governed by genetic mechanisms that are homologous and deeply conserved across a wide range of species.

<span class="mw-page-title-main">Holozoa</span> Clade containing animals and some protists

Holozoa is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa and the protists Choanoflagellata, Filasterea, Pluriformea and Ichthyosporea. Along with fungi and some other groups, Holozoa is part of the Opisthokonta, a supergroup of eukaryotes. Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic.

<i>Volvox carteri</i> Species of alga

Volvox carteri is a species of colonial green algae in the order Volvocales. The V. carteri life cycle includes a sexual phase and an asexual phase. V. carteri forms small spherical colonies, or coenobia, of 2000–6000 Chlamydomonas-type somatic cells and 12–16 large, potentially immortal reproductive cells called gonidia. While vegetative, male and female colonies are indistinguishable; however, in the sexual phase, females produce 35-45 eggs and males produce up to 50 sperm packets with 64 or 128 sperm each.

<span class="mw-page-title-main">Allorecognition</span>

Allorecognition is the ability of an individual organism to distinguish its own tissues from those of another. It manifests itself in the recognition of antigens expressed on the surface of cells of non-self origin. Allorecognition has been described in nearly all multicellular phyla.

A somatic mutation is a change in the DNA sequence of a somatic cell of a multicellular organism with dedicated reproductive cells; that is, any mutation that occurs in a cell other than a gamete, germ cell, or gametocyte. Unlike germline mutations, which can be passed on to the descendants of an organism, somatic mutations are not usually transmitted to descendants. This distinction is blurred in plants, which lack a dedicated germline, and in those animals that can reproduce asexually through mechanisms such as budding, as in members of the cnidarian genus Hydra.

References

  1. Grosberg, Richard K.; Strathmann, Richard R. (2007). "The Evolution of Multicellularity: A Minor Major Transition?". Annual Review of Ecology, Evolution, and Systematics. 38: 621–654. doi:10.1146/annurev.ecolsys.36.102403.114735.
  2. Atala, Anthony; Lanza, Robert (2013). Handbook of Stem Cells. Academic Press. ISBN   9780123859433.
  3. Niklas, Karl J (Jan 2014). "The evolutionary-developmental origins of multicellularity". American Journal of Botany. 101 (1): 6–25. doi:10.3732/ajb.1300314. PMID   24363320.
  4. Bernardes, Joana P.; John, Uwe; Woltermann, Noemi; Valiadi, Martha; Hermann, Ruben J.; Becks, Lutz (2021-07-09). "The evolution of convex trade-offs enables the transition towards multicellularity". Nature Communications. 12 (1): 4222. Bibcode:2021NatCo..12.4222B. doi:10.1038/s41467-021-24503-z. ISSN   2041-1723. PMC   8270964 . PMID   34244514.
  5. 1 2 Chase, Jonathan M. (2014-05-13). "A Fool to Do Your Dirty Work?". PLOS Biology. 12 (5): e1001859. doi: 10.1371/journal.pbio.1001859 . ISSN   1544-9173. PMC   4019462 . PMID   24823481.
  6. Goldsby, Heather J.; Knoester, David B.; Ofria, Charles; Kerr, Benjamin (2014-05-13). Keller, Laurent (ed.). "The Evolutionary Origin of Somatic Cells under the Dirty Work Hypothesis". PLOS Biology. 12 (5): e1001858. doi: 10.1371/journal.pbio.1001858 . ISSN   1545-7885. PMC   4019463 . PMID   24823361.
  7. Gilbert, Scott F. (2000). "Multicellularity: The Evolution of Differentiation". Developmental Biology. 6th Edition.
  8. Hallmann, Armin (June 2011). "Evolution of reproductive development in the volvocine algae". Sexual Plant Reproduction. 24 (2): 97–112. doi:10.1007/s00497-010-0158-4. ISSN   0934-0882. PMC   3098969 . PMID   21174128.