Cell wall

By | July 25, 2021

Cell wall

The cells of plants and algae, fungi and most chromalveolates have a cell wall, a layer outside the cell membrane, providing the cell with structural support, protection, and a filtering mechanism. The cell wall also prevents over-expansion when water enters the cell.

The major polysaccharides making up the primary cell wall of land plants are cellulose, hemicellulose, and pectin. The cellulose microfibrils are linked via hemicellulosic tethers to form the cellulose-hemicellulose network, which is embedded in the pectin matrix. The most common hemicellulose in the primary cell wall is xyloglucan.

Differences among eukaryotic cells:

There are many different types of eukaryotic cells, though animals and plants are the most familiar eukaryotes, and thus provide an excellent starting point for understanding eukaryotic structure. Fungi and many protists have some substantial differences, however.

Animal cell:

Structure of a typical animal cell:

Structure of a typical plant cell:

All animals are eukaryotic. Animal cells are distinct from those of other eukaryotes, most notably plants, as they lack cell walls and chloroplasts and have smaller vacuoles. Due to the lack of a cell wall, animal cells can transform into a variety of shapes. A phagocytic cell can even engulf other structures.

 Plant cell

Plant cells are quite different from the cells of the other eukaryotic organisms. Their distinctive features are:

  • A large central vacuole (enclosed by a membrane, the tonoplast), which maintains the cell’s turgor and controls movement of molecules between the cytosol and sap
  • A primary cell wall containing cellulose, hemicellulose and pectin, deposited by the protoplast on the outside of the cell membrane; this contrasts with the cell walls of fungi, which contain chitin, and the cell envelopes of prokaryotes, in which peptidoglycans are the main structural molecules
  • The plasmodesmata, pores in the cell wall that link adjacent cells and allow plant cells to communicate with adjacent cells. Animals have a different but functionally analogous system of gap junctions between adjacent cells.
  • Plastids, especially chloroplasts, organelles that contain chlorophyll, the pigment that gives plants their green color and allows them to perform photosynthesis
  • Bryophytes and seedless vascular plants only have flagellae and centrioles in the sperm cells. Sperm of cycads and Ginkgo are large, complex cells that swim with hundreds to thousands of flagellae.
  • Conifers (Pinophyta) and flowering plants (Angiospermae) lack the flagellae and centrioles that are present in animal cells.

Fungal cell:

Fungal Hyphae cells: 1 – hyphal wall, 2 – septum, 3 – mitochondrion, 4 – vacuole, 5 – ergosterol crystal, 6 – ribosome, 7 – nucleus, 8 – endoplasmic reticulum, 9 – lipid body, 10 – plasma membrane, 11 – spitzenkörper, 12 – Golgi apparatus

The cells of fungi are most similar to animal cells, with the following exceptions:

  • A cell wall that contains chitin
  • Less compartmentation between cells; the hyphae of higher fungi have porous partitions called septa, which allow the passage of cytoplasm, organelles, and, sometimes, nuclei. Primitive fungi have few or no septa, so each organism is essentially a giant multinucleate supercell; these fungi are described as coenocytic.
  • Only the most primitive fungi, chytrids, have flagella.

Other eukaryotic cells:

Some groups of eukaryotes have unique organelles, such as the cyanelles (unusual chloroplasts) of the glaucophytes, the haptonema of the haptophytes, or the ejectosomes of the cryptomonads. Other structures, such as pseudopodia, are found in various eukaryote groups in different forms, such as the lobose amoebozoans or the reticulose foraminiferans.

Reproduction:

This diagram illustrates the twofold cost of sex. If each individual were to contribute to the same number of offspring (two), (a) the sexual population remains the same size each generation, where the (b) asexual population doubles in size each generation.

Cell division generally takes place asexually by mitosis, a process that allows each daughter nucleus to receive one copy of each chromosome. Most eukaryotes also have a life cycle that involves sexual reproduction, alternating between a haploid phase, where only one copy of each chromosome is present in each cell and a diploid phase, wherein two copies of each chromosome are present in each cell. The diploid phase is formed by fusion of two haploid gametes to form a zygote, which may divide by mitosis or undergo chromosome reduction by meiosis. There is considerable variation in this pattern. Animals have no multicellular haploid phase, but each plant generation can consist of haploid and diploid multicellular phases.

Eukaryotes have a smaller surface area to volume ratio than prokaryotes, and thus have lower metabolic rates and longer generation times.

The evolution of sexual reproduction may be a primordial and fundamental characteristic of eukaryotes. Based on a phylogenetic analysis, Dacks and Roger proposed that facultative sex was present in the common ancestor of all eukaryotes. A core set of genes that function in meiosis is present in both Trichomonas vaginalis and Giardia intestinalis, two organisms previously thought to be asexual. Since these two species are descendants of lineages that diverged early from the eukaryotic evolutionary tree, it was inferred that core meiotic genes, and hence sex, were likely present in a common ancestor of all eukaryotes. Eukaryotic species once thought to be asexual, such as parasitic protozoa of the genus Leishmania, have been shown to have a sexual cycle. Also, evidence now indicates that amoebae, previously regarded as asexual, are anciently sexual and that the majority of present-day asexual groups likely arose recently and independently.

Classification:

Phylogenetic and symbiogenetic tree of living organisms, showing a view of the origins of eukaryotes & prokaryotes

One hypothesis of eukaryotic relationships – the Opisthokonta group includes both animals (Metazoa) and fungi, plants (Plantae) are placed in Archaeplastida.

A pie chart of described eukaryote species (except for Excavata), together with a tree showing possible relationships between the groups

In antiquity, the two lineages of animals and plants were recognized. They were given the taxonomic rank of Kingdom by Linnaeus. Though he included the fungi with plants with some reservations, it was later realized that they are quite distinct and warrant a separate kingdom, the composition of which was not entirely clear until the 1980s. The various single-cell eukaryotes were originally placed with plants or animals when they became known. In 1818, the German biologist Georg A. Goldfuss coined the word protozoa to refer to organisms such as ciliates, and this group was expanded until it encompassed all single-celled eukaryotes, and given their own kingdom, the Protista, by Ernst Haeckel in 1866. The eukaryotes thus came to be composed of four kingdoms:

  • Kingdom Protista
  • Kingdom Plantae
  • Kingdom Fungi
  • Kingdom Animalia

The protists were understood to be “primitive forms”, and thus an evolutionary grade, united by their primitive unicellular nature. The disentanglement of the deep splits in the tree of life only really started with DNA sequencing, leading to a system of domains rather than kingdoms as top level rank being put forward by Carl Woese, uniting all the eukaryote kingdoms under the eukaryote domain. At the same time, work on the protist tree intensified, and is still actively going on today. Several alternative classifications have been forwarded, though there is no consensus in the field.

Eukaryotes are a clade usually assessed to be sister to Heimdallarchaeota in the Asgard grouping in the Archaea. The basal groupings are the Opimoda, Diphoda, the Discoba, and the Loukozoa. The Eukaryote root is usually assessed to be near or even in Discoba.

A classification produced in 2005 for the International Society of Protistologists, which reflected the consensus of the time, divided the eukaryotes into six supposedly monophyletic ‘supergroups’. However, in the same year (2005), doubts were expressed as to whether some of these supergroups were monophyletic, particularly the Chromalveolata, and a review in 2006 noted the lack of evidence for several of the supposed six supergroups. A revised classification in 2012 recognizes five supergroups.

Archaeplastida (or Primoplantae) Land plants, green algae, red algae, and glaucophytes
SAR supergroup Stramenopiles (brown algae, diatoms, etc.), Alveolata, and Rhizaria (Foraminifera, Radiolaria, and various other amoeboid protozoa)
Excavata Various flagellate protozoa
Amoebozoa Most lobose amoeboids and slime molds
Opisthokonta Animals, fungi, choanoflagellates, etc.

There are also smaller groups of eukaryotes whose position is uncertain or seems to fall outside the major groups – in particular, Haptophyta, Cryptophyta, Centrohelida, Telonemia, Picozoa, Apusomonadida, Ancyromonadida, Breviatea, and the genus Collodictyon. Overall, it seems that, although progress has been made, there are still very significant uncertainties in the evolutionary history and classification of eukaryotes. As Roger & Simpson said in 2009 “with the current pace of change in our understanding of the eukaryote tree of life, we should proceed with caution.”

In an article published in Nature Microbiology in April 2016 the authors, “reinforced once again that the life we see around us – plants, animals, humans and other so-called eukaryotes – represent a tiny percentage of the world’s biodiversity.” They classified eukaryote “based on the inheritance of their information systems as opposed to lipid or other cellular structures.” Jillian F. Banfield of the University of California, Berkeley and fellow scientists used a super computer to generate a diagram of a new tree of life based on DNA from 3000 species including 2,072 known species and 1,011 newly reported microbial organisms, whose DNA they had gathered from diverse environments. As the capacity to sequence DNA became easier, Banfield and team were able to do metagenomic sequencing – “sequencing whole communities of organisms at once and picking out the individual groups based on their genes alone

Phylogeny:

The rRNA trees constructed during the 1980s and 1990s left most eukaryotes in an unresolved “crown” group (not technically a true crown), which was usually divided by the form of the mitochondrial cristae; see crown eukaryotes. The few groups that lack mitochondria branched separately, and so the absence was believed to be primitive; but this is now considered an artifact of long-branch attraction, and they are known to have lost them secondarily.

As of 2011, there is widespread agreement that the Rhizaria belong with the Stramenopiles and the Alveolata, in a clade dubbed the SAR supergroup, so that Rhizaria is not one of the main eukaryote groups; also that the Amoebozoa and Opisthokonta are each monophyletic and form a clade, often called the unikonts. Beyond this, there does not appear to be a consensus.

It has been estimated that there may be 75 distinct lineages of eukaryotes. Most of these lineages are protists.

The known eukaryote genome sizes vary from 8.2 megabases (Mb) in Babesia bovis to 112,000–220,050 Mb in the dinoflagellate Prorocentrum micans, showing that the genome of the ancestral eukaryote has undergone considerable variation during its evolution. The last common ancestor of all eukaryotes is believed to have been a phagotrophic protist with a nucleus, at least one centriole and cilium, facultatively aerobic mitochondria, sex (meiosis and syngamy), a dormant cyst with a cell wall of chitin and/or cellulose and peroxisomes.Later endosymbiosis led to the spread of plastids in some lineages.

Discussion of Phylogenetic Relationships:

Our understanding of eukaryotic relationships has been transformed by the use of molecular data to reconstruct phylogenies (Sogin et al., 1986). Prior to that, the diversity of microbial eukaryotes was vastly underestimated, and the relationships between them and multicellular eukaryotes were difficult to resolve (Taylor, 1978). Early molecular phylogenies based on small subunit ribosomal RNA (SSU rRNA) gene sequences suggested a ladder of basal lineages topped by a ‘crown’ composed of multicellular groups (animals, plants, and fungi) together with a subset of the purely microbial lineages (Sogin, 1989). A great number of the relationships revealed by SSU rRNA phylogeny have stood the test of time, but subsequent analyses based on protein coding genes and more recently very large datasets composed of hundreds of protein coding genes have led to a revision of the overall structure of the tree. The current view of eukaryotic phylogeny is of a small number of large ‘supergroups’, each comprising a spectacular diversity of structures, nutritional modes, and behaviours (Adl et al., 2005; Keeling, 2004; Keeling et al., 2005; Simpson and Roger, 2002). Some of these supergroup hypotheses are well supported, while others remain the subject of vigorous debate (see (Keeling et al., 2005) for a discussion of evidence).  Furthermore the relationships between supergroups are poorly understood. Below we summarise the main members of each supergroup, the evidence for its monophyly, and emerging hypotheses for inter-supergroup relationships.

Archaeplastida (Plantae):

The Archaeplastida, or Plantae, comprises glaucophytes, red algae, green algae and plants. They are united by the possession of a plastid derived from primary endosymbiosis (see Symbiosis section). There has long been strong support for the monophyly of plastids in Archaeplastida based on molecular phylogeny and also plastid genome structure (Turner, 1997; Turner et al., 1999), and molecular phylogenies based on large numbers of protein coding genes have more recently demonstrated the monophyly of the nuclear/cytosolic lineage as well (Burki et al., 2008; Moreira et al., 2000; Reyes-Prieto et al., 2007).

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