An Introduction to Molecular Biotechnology. Группа авторов

endocytosis polar molecules can also enter the cell, which otherwise would not be taken up via diffusion or carriers.

An important variation of endocytosis is receptor‐mediated endocytosis. This is how lipoproteins such as low‐density lipoprotein(LDL) particles, which are loaded with cholesterol ester in blood, are recognized and bound by LDL receptors of the target cell (Figure 5.10). After binding, the clathrin‐coated endocytosis vesicle buds off and migrates via the endosomes to the lysosomes. There the receptors with exocytotic vesicles are returned to the cytoplasmic membrane, while the lipoproteins are degraded in the lysosome. The cholesterol ester is also cleaved by an esterase. Cholesterol is then available to the cell for synthesis or as a membrane lipid. Patients with defective LDL receptor genes (prevalence of 1 in 500) have an increased risk of myocardial infarction, as higher levels of cholesterol lead to arteriosclerosis.

Figure 5.10 Schematic progression of receptor‐mediated endocytosis of LDL.

      Source: From Voet et al. (2016).

References

      1 Nigg, E.A. (1997). Nucleocytoplasmic transport: signals, mechanisms and regulation. Nature 386: 779–787.

      2 Voet, D., Voet, J.G., and Pratt, C.W. (2016). Fundamentals of Biochemistry, Live at the Molecular Level, 5e. Holboken, NJ: Wiley.

Further Reading

      1 Alberts, B., Johnson, A., Lewis, L. et al. (2015). Molecular Biology of the Cell, 6e. New York: Garland Science.

      2 Alberts, B., Bray, D., Hopkin, K. et al. (2019). Essential Cell Biology, 5e. New York: Garland Science.

      3 Krebs, J., Goldstein, E.S., and Kilpatrick, S.T. (2018). Lewin's Genes XII. Burlington: Jones & Bartlett Learning.

6 Evolution and Diversity of Organisms

       Michael Wink

       Heidelberg University, Institute of Pharmacy and Molecular Biotechnology (IPMB), Im Neuenheimer Feld 329, 69120 Heidelberg, Germany

Molecular cell biology focuses more and more on model organisms. Examples are, among others, the Gram‐negative bacteria Escherichia coli, the brewer's yeast (Saccharomyces cerevisiae), the nematode worm (Caenorhabditis elegans), the fruit fly (Drosophila melanogaster), the common mouse (Mus musculus), the zebra fish (Danio rerio), the human (Homo sapiens), and wall cress (Arabidopsis thaliana) as a representative of higher plants. Extensive knowledge about molecular and cellular biology was gained from these model organisms. As these organisms share a common evolutionary history, it is assumed that basic characteristics found in one model organism are also valid for all other organisms. This can, but must not always, be true. In some cases nature has also found different solutions to the same problem (convergent evolution). Nevertheless, we see an astonishing degree of similarity in fundamental mechanisms and structures.

      However, there is also an astonishing diversity at the organismal level. In the future of biotechnology, we must not ignore the diversity of organisms, with presently known 2 million species (perhaps over 10 million species exist) and complex adaptations. Many of them offer evolutionarily derived solutions for problems that are of great interest for biotechnological purposes or applications.

6.1 Prokaryotes

Figure 6.1 shows a tree of life reconstructed over nucleotide sequences of 31 genes from 191 species whose genomes were completely sequenced (Ciccarelli et al. 2006). This tree illustrates the lines of development for the different kingdoms. Within prokaryotes, two large domains are recognizable: the Bacteria (or simply bacteria) and the Archaea (or archaebacteria). Important biochemical differences were already summarized in Table 1.1.

Figure 6.1 A phylogenetic tree of life, showing the relationship between species whose genomes had been sequenced as of 2006. The very center represents the last universal ancestor of all life on earth. The different colors represent the three domains of life: pink represents eukaryota (animals, plants, and fungi), blue represents bacteria, and green represents archaea. Note the presence of Homo sapiens (humans) second from the rightmost edge of the pink segment. The light and dark bands along the edge correspond to clades: the rightmost light red band is Metazoa, with dark red Ascomycota to its left, and light blue Firmicutes to its right.

      Source: https://commons.wikimedia.org/wiki/File:Tree_of_life_SVG.svg..

6.2 Eukaryotes

The evolution of the ancestor of a eukaryotic cell and the uptake of bacteria (endosymbiotic origin of mitochondria and chloroplasts) were a key innovation of early evolution. While the incorporation of mitochondria only occurred once in evolution, there is reasonable evidence for the assumption that the incorporation of cyanobacteria (leading to chloroplasts) occurred many times (especially within different groups of algae).

      There are large differences in cellular structure and function between prokaryotes and eukaryotes. Table 1.1 summarized the important characteristics. The eukaryotic cell is distinctly further developed (Figure 1.2) and is able to carry out different processes at the same time in a single cell. This required the development of separated reaction spaces – cellular compartments (Table 1.2) – in the early stages of evolution.

A simplified overview of the origin of organisms is shown in Figures 1.1 and 6.1. Due to lack of space, it is not possible to go into more detail for the different organisms in the specific individual domains of the living kingdoms. To give biotechnologists a quick orientation about which organism they are focusing on and where these organisms stand in the tree of life, a short systematic synopsis of the organisms is put together in the following. For simplicity, only the large groups of protists (Table 6.1; Figure 6.2), plants (Table 6.2; Figure