Rise of The Eukaryotes

Rise of the Eukaryotes: The transition from prokaryotic life.
It is generally believed that prokaryotic cells predate eukaryotic cells, due to the complete absence
in prokaryotes of several cell structures that are present in eukaryotes. The transition from
prokaryotes to eukaryotes is a major evolutionary event which is thought to be the result of
endosymbiosis by a eubacterial prokaryotic cell, of an archaebacterium to create an altogether new
cellular domain: that of the eukaryotes (Stearns and Hoekstra, 2005). The transition was marked by
the acquisition of the cytoskeleton (Horiike et al., 2001), cell organelles such as mitochondria and
chloroplasts (Martin et al., 2002), and the addition of a nuclear membrane to protect genetic
material. The transition enabled several other major evolutionary events, such as the organisation of
cells to form multi-cellular organisms, and therefore occupies a central role in the current form of
evolutionary theory (Smith and Szathmary, 2005).
Many different theories have been posited for the transition between prokaryotes and eukaryotes,
especially for the acquisition of various cell organelles and the nucleus, however that which remains
most convincing is the theory of endosymbiosis, first conjectured by Lynn Margulis in 1970. The
theory states that the development of eukaryotes began with eubacteria, which have a cytoskeleton
instead of a cell wall and feed via the process of phagocytosis, which involves taking smaller
organisms inside the eubacterial cell and digesting them. It is thought that at some point the
phagocytotic process failed, leaving functional archaebacterium inside the eubacterial cell, where it
did not impact negatively on the fitness of the organism, survived and reproduced in an autonomous
fashion (although presumably there were some cases where this kind of mutually beneficial
relationship did not unfold).
The nuclear membrane is thought to have originated when parts of the host genome were absorbed
into the archea cell. This collection of the genetic material would have served to increase the fitness
of the host by protecting the DNA from damage, either by ultra violet light, or exposure to
metabolic by-products in the cytoplasm, and also by cetralising the copying process, which would
probably have made reproduction faster.
The eukaryotic genome contains traces to support this theory of its origin: the somewhat quixotic
structure, as revealed by 16s rRNA, shows similarity in the genetic coding for some proteins,
particularly histones, as well as highlighting many similarities to both the archea and eubacterial
cellular domains (Horiike et al., 2001).
Variants on this theory can also apply to the acquisition of other organelles such as mitochondria
and chloroplasts (Bogorad, 2007)(Martin et al., 2002). Assuming a similar situation with an
anaerobic proto-eukaryote (with or without a nucleus) a bacterium which escaped digestion would
be able to provide its host with a range of beneficial metabolites and in turn receive a steady supply
of resources as well as protection from predation(Spring, 2003)(Finlay and Esteban, 2009).
Traditional theories suggest that the acquisition of chloroplasts and mitochondria were independent
endosymbiotic events with cyanobacterium (Martin et al., 2002)(Keeling, 2004) and protobacterium
(Emelyanov, 2003) respectively. A secondary loss of respiratory metabolism in chloroplasts and
photosynthetic metabolism in mitochondria occurred due to selection pressures against the
unnecessary metabolism in question (Stearns and Hoekstra, 2005). Evidence suggests that since this
time genetic information for both chloroplasts and mitochondria have been steadily migrating to the
nucleus in the same way that the nucleus acquisition theory outlines (Bogorad, 2007). It is
interesting to note however that although the nucleus now holds the entire genome for both original
host and symbiont, the nuclear genome does not hold the genes pertaining to metabolic the
processes within the organelles. This is more than likely due to the time sensitive nature of the
metabolism taking place within these organelles. Having the genes on site mean having a much
quicker response to feedback and expression of these genes whilst managing the processes these
organelles handle (Martin et al., 2002)(Stearns and Hoekstra, 2005).
Currently the most accepted theory for the genesis and development of the eukaryotic cytoskeleton
stems from the prokaryotic cell wall. The cell wall provides mechanical support for the cell
preventing it from bursting from turgid pressure in aquatic environments (due to the potential
isotonic imbalance experienced between the cell interior and the environment) as well as providing
protection from other mechanical forces. The problem encountered by a prokaryotic cell utilising
such a structure is that it is restrictive, that is to say that transfer of materials in and out of the cell
may only be as large as the pores in the cell wall (Stearns and Hoekstra, 2005).
The development of a cytoskeleton enabled similar mechanical support to be provided through an

intercellular network of actin microfilaments and microtubules, and also brought a host of other
adaptive advantages. It brought about intracellular transport of resources (and later organelles), as
well as motility of the cell itself through its environment not unlike amoeba and other protozoa do
today with pseudopodia. The absence of the restrictive cell wall pores would also open the door to
the predation of smaller organisms through phagocytosis (Stearns and Hoekstra, 2005).
Cytoskeletons also helped in the evolution of faster and more efficient cell division (Faguy and
Doolittle, 1998). Finally the major restriction lifted was that of the genome size an organism could
support. In traditional prokaryotes a single chromosome is anchored at two points to the cell wall.
With the loss of a cell wall and a dynamic cytoskeleton in its place, eukaryotes now had the ability
to support multiple chromosomes. This paved the way for much larger genomes and consequently
code for a greater range of complexity and adaptive advantages. In addition multiple point DNA
replication was now possible, meaning that the time taken to copy a large genome no longer
represented an upper limit to genome size like it had previously due to the short generational time
of these organisms. In this it could be seen that although a cytoskeleton was not able to provide all
of the strength that a cell wall provided against environmental forces, the lower resource cost
coupled with the advantages listed above allowed the cytoskeleton endowed cell to be much more
versatile and adaptive.
These changes resulted in a cell divided into sections by purpose, from anabolic and catabolic
metabolism within the chloroplasts and mitochondria to gene expression and regulation within the
nucleus. This level of organisation and compartmentalisation coupled with the dynamic internal
structure of the cytoskeleton produced a cell capable of more complicated feats and internal
regulation than its prokaryotic precursors making eukaryotes a base for rapid expansions in
evolutionary history.
Bibliography
Finlay, B.J. and Esteban, G.F. (2009) 'Can biological complexity be rationalised?', BioScience, pp.
333 - 340.
Hartman, H. and Fedorov, A. (2002) 'The origin of the eukaryotic cell: A genomic investigation',
Proceedings of the National Academy of Sciences, pp. 1420 - 1425.
Horiike, T., Hamada, K., Kanaya, S. and Shinozawa, T. (2001) 'Origin of eukaryotic cell nuclei by
symbiosis of Archaea in Bacteria is revealed by homology-hit analysis.', Nature Cell Biology, pp.
210 - 214.
Keeling, P.J. (2004) 'Diversity and evolutionary history of plastids and their hosts', American
Journal of Botany, pp. 1481 - 1493.
Martin, W., Rujan, T., Richly, E., Hansen, A., Coronelsen, S., Lins, T., Leister, D., Stoebe, B.,
Hasegawa, M. and Penny, D. (2002) 'Evolutionary analysis of Arabidopsis, cyanobacterial, and
chloroplast genomes reveals plastid phylogeny and thousands of cyanobacterial genes in the nucleus
', Proceedings of the National Academy of Sciences of the United States of America, pp. 12246 -
12251.
Maynard Smith, J. and Szathmary, E. (1997) 'From Replicators to Reproducers The first major
transitions leading to life', Journal of Theoretical Biology, pp. 555 - 571.
Maynard Smith, J. and Szathmary, E. (2005) The Major Transitions in Evolution, Oxford
University Press.
Spring, J. (2003) 'Major transitions in evolution by genome fusions: from prokaryotes to

eukaryotes, metazoans, bilaterians and vertebrates', Journal of Structural and Functional Genomics,
pp. 19 - 25.
Stearns, S.C. and Hoekstra, R.F. (2005) Evolution, an Introduction, Oxford: Oxford University
Press.

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Rise of the Eukaryotes: The transition from prokaryotic life.

The development of a cytoskeleton enabled similar mechanical support to be provided through an

Spring, J. (2003) 'Major transitions in evolution by genome fusions: from prokaryotes to

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