Many people understand the functions of the intracellular organelles, the mitochondria and

the nucleus in eukaryotic cells, but few know of the origins of these organelles. This paper

explores the origins of the mitochondria and the nucleus by looking at how these organelles

evolved into their modern-day counterparts by outcompeting their neighbours and what they

provide to the cell that other organelles cannot.

The mitochondria have a few functions that they provide to the cell that make it unique in its

purpose where its main function is aerobic respiration/cellular respiration. There is common

ground for the origin of the mitochondria is that there was an ancestral endosymbiont that

had collided with a prokaryotic cell to form a eukaryotic cell. Most people don't know the

origin of the mitochondria. There are various competing hypotheses on the origins of the

mitochondria and several hypotheses and proposed models (figure 1) for the origins of the

mitochondria supported by strong evidence of a host and a “guest”1. Where there were

multiple possibilities of the type of host and the “guest” inside the host is how the hypotheses

had differed as shown in Table 1 of the beneficial interaction and costly interactions between

the “guest” and the host. Table one shows the interactions between the different types of

posts and the ecological relationship with each of the hypotheses.

There are generally 2 proposed pathways of the eukaryotic cell and the mitochondria where

the mitochondria come later then the eukaryotic cell develops on its own beforehand and the

mitochondria are brought into the cell or where the mitochondria develop first and the

eukaryotic cell come later2. However, these classifications are quite limited due to their

simplistic nature when tackling a topic as complex as the origin of the mitochondria. Where

there are unavoidable questions that arise such as what was the origin metabolism of the

host? the stability of the partnership of the host cell and the mitochondria? and the early

selective advantage that the partnership granted over other organelles? 3 the different
hypotheses that try to answer the different unavoidable questions that are part of trying to

find the origin of the mitochondria whereas some hypothesize answered more of the

unavoidable questions more solidly.

Origin and diversification

Where a pre-mitochondrial-alphaproteobacterium would have done endosymbiosis within a

cell despite how it was unclear how the endosymbiosis was achieved in the end4. Where the

endosymbiont had lost its autonomy when it became integrated with the host5. After

integration, the endosymbiont slowly changed into the modern-day mitochondria through a

long multistep process that ended up with the mitochondria having a more simplistic genome

and no autonomy compared to their bacterium relatives6. Where the host could insert the

protein machinery and pathways needed to gain the endosymbiont ATP supply to be used in

the cell and to gain control over the endosymbiont envelope7. The endosymbiont had lost its

original identity over time due to the rewriting and simplification of its genome within the

host8. The common ancestor had about 3000 genes and the alphaproteobacterium had a

range between 800-8000 complete genes compared to the genes of the mitochondria which

were 699. The reduction of genes happened in the pre-mitochondrial evolution that made it

so that the endosymbiont could not replicate outside of the host cell10. The mitochondrial

genome could be reduced even further and import the host's genome and mitochondria and

EGT that could be reduced further along the eurkotyic tree11.

Figure 1: The 8 different hypotheses how how the endosymbiont had gone into the cell.

Nucleus origin
The chimeric eukaryote: Origin of the

nucleus from the karyomastigont in

amitochondriate protists
Lynn Margulis,* Michael F. Dolan,*† and Ricardo Guerrero‡

The nucleus has a few functions which are the storage of genetic information DNA/RNA and

regulating activities of the cell like metabolism and when to do mitosis and divide the cell.

The functions of the nucleus are well known to people but the origins of the nucleus are

unknown to the majority of people. The origins of the nucleus has no solid answer but there

are several competing hypothesis about the origin of the nucleus. The hypothesis that will be

talked about in this paper is that the nucleus as a testable model is a chimera of more than

one prokaryote1. Where the first anaerobic eukaryotes had descended from two prokaryotic

lineages2. The enzymes of protein synthesis in eurkorayes come from archaea bacteria in

the motility system, many soluble heat shocks and other proteins from eubacteria3 and use

protist data to make the testable model and recreate the fusion to allow for the formation of

the nucleus4.

A study was done by Gupta on protein sequences where the conclusion was found that

eukaryotic cells had received major contributions from arecia bacteria (Gupta, ). Where

ancestral eukaryotic cells aren't direct descendants from archaebacteria but rather a

chimaera of the integration and fusion of the genomes of the archaebacteria and

gram-negative bacteria5. The archaebacteria haven't been identified but the archaebacteria

could have come from a Thermoplasma acidophilum-like thermoacidophilic (eocyte)

prokaryote6. The conditions for this archaebacteria ancestor had lived in acidic, warm, and
sporadically sulphurous water that had used either elemental sulphur or 5% oxygen (

generating H2O) as an electron acceptor.7 The ancient archaebacteria survived this acid

hydrolysis environmental conditions due to nucleosome histone protein coating of its DNA

and protein synthesis like actin8. The second member of the consortium was anaerobic

which interacted with sulphur and sulphate to reduce these conditions and produce

hydrogen sulfide9. Degradation of carbohydrates and oxidation of sulphide to elemental

sulphur by the carbon-rich eubacteria fermentation products and their electron acceptors for

when the eubacteria and the archaebacteria had fused to form the consortium had the

likelihood of increasing the amount of carbon it had and that likely helped with the formation

of the chimaera (figure 2). The chimaera is slightly different from the geothermically

important “Thiodendron”10.

Figure 2: the origin of chimeric eukaryotes with karyomastigont from a moltile sulphur

bacteria consortium.

The “thioderion” stage refers to the extant bacterial consortium. Where the partners had

become the chimeric predecessors to archaea protists 11. Morphologically the

membrane-bound nucleus is the chimaera genetic system made manifest12. A study that

had been made with marine microbial mats showed prevalent bacterial consortia in several

geologically isolated locations133 where they all contained "Thiodendron latens” or similar

bacteria. Samples were taken in water with just below average oxygen sulphide levels in

anoxic water. Where additional lab work was done to abolish the thiodendron genus due to

having sulphur syntrophy14. A stable ectosymbiont bacterial association is grown as a

consortmation that gets its carbon from heterotrophic carbon dioxide fixation of

carbohydrates like starch, and cellobiose and exists in stable temperatures between 4 and

32oC. The thioderion is engulfed by the consortium bacteria and appears as gelatinous
bluish-white colonies in a smiley matrix. There is a dominant partner which was a distinct

strain of pleomorphic spirochetes where they had varied in their shape, size, and distinct

features15. Then the non-dominant partner of the consortium is a small morphologically

stable vibrioid, Desulfobacter sp that needs organic carbon to break down through

spirochetal carbohydrate degradation and fermentation and generate its food and energy 16.

Desulfobacter that reduce sulphide and sulphur are also present but much rare in this study

17.

When “pure cultures” had survived low oxygen levels they had shown a complex history of

fibroids, spheroids, threads and helices connected to “Thiodendron latens” when observed

by B. V. Perfil'ev in 1969 18. Where these histories are due to environmental pressures like if

the redox potential is changed by addition or removal of a sulphide compound19. The growth

of the two partners depended upon the bacterial development patterns of the Thiodendron.

Where the oxygen level is lowered to maintain the syntropy. Where sulphur

oxidation-reduction and oxygen removal from oxygen-sensitive enzymes could have been

internalised by the chimaera and retained by protists' descendants as the development

continues20.

The two unlike prokaryotes together produced a consistent protein package. Where this part

of the nucleus origin is traceable to the morphological legacy of the chimera the

karyomastigont21. The attached swimmer is the precursor of the mitotic microtubule system.

The swimmers' attached structures hypertrophied as they would typically in extant motility

symbioses22. Where the swimmer attachment systems of the archaebacterium-eubacterium

then become the karyomastigont23. The karyomastigont unites the partner's DNA into a

membrane-bound package assuring the of the chimera24. Then the chimera's DNA

reorignized itself and the protein-based motility system segregated the remaining chimera
DNA which is retained by amitochondriate protists and later their mitochondrial descendants.

Unattached nuclei also evolved many times to dissociate from the rest of the

karyomastigont. Leading to the karyomastigont being the first cellular microtubule

organisation center25.

The karyomastigont had preceded nuclei

The term “karyomastigont” was coined by Janicki which refers to the organ-cellular system

observed in certain protists consisting of two parts the “cell whip” like eukaryotic flagellum

and the “nuclear connecter” connecting to a nucleus 26. The term was used after the results

from Janicki's work on termite intestines where karyomasigonts dominate highly motile

trichomonads symibonts27.

The karyomastigont is an ancestral feature of eukaryotes and is found in “early branching”

protists28. Archaeoprotisits are heterotrophic unicells that exist in anoxic habitats that all

lack mitochondria29. These archaeologists could have either beared karyomastigont or had

karyomastigont deprived of simple morphological steps in organelle reproduction30 (Table 1

appendix 1). During the evolution of the protists the nuclei would be severed from the

karyomastigont and akaryomasigonts were generated31. Nuclei unattached were able to

temporarily were able to proliferate and have central positions within a cell 32. which helps

generate larger and faster swimming cells in the same evolutionary step 33. The

karyomastigonts are the cytoskeleton for almost all archaeologist lineages (three classes:

Archamoeba, Metamonads, and Parabasalia)34(figure 2).

Figure 3: biological pathways for chimeric eukaryotes into chromotrites.

In trichomonads, the karyomastigonts also include a Golgi complex and coordinate the

placement of hydrogenosomes (membrane-bound bacterial cell inclusion that generates

hydrogens). The karyomasigonts are reproduced as a unit structure 35. Typically four

attached kinesomes with rolled sheets of microtubules so they retained their morphological

relationships as they reproduce36. Kinetosomes reproduce first then the nucleus will divide,
and the the two kinetosome groups separate at the poles of a thin microtubule spindle37.

Then kinetosomes with other associated structures formed into one of the two

karyomastigonts. The other karyomasigont produced components the first one lacked like

the Golgi complex38. Nuclear α-proteobacterial could have originated from lost or

degenerated genes from the mitochondria in at least two species of archaea protists 39

(Giardia lamblia, Trichomonas vaginalis). The mitochondria were never lost in the ancestors

and were more likely lost in anaerobic protists40. Eubacterial genes are acquired in

amitochondriate protists from proteobacterial symbionts other than those of the

mitochondrial lineage41. Karyomastigont detached from nuclei independently from before

and after the acquisition of the mitochondria42.