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How Did Complex Living Cells Evolve?

Paul Kotschy

25 August 2023

phacus-gigas.jpg
The unicellular protozoan Phacus gigas packed with endosymbiont algae cells living inside P. gigas's cyctoplasm (630×).
kidney-cell-mitochondrion.jpg
Mitochondria (false-coloured red) in a cell from a tubule in the kidney cortex. The nucleus of the cell has a prominent nucleolus (false-coloured dark brown).

The structural and functional complexity1 2 found in and between living cells is remarkable. Whence comes this complexity? It might be tempting to appeal to the notion of irreducible complexity, and thereby to assert that living cells could not have evolved, but instead were designed and constituted by some non-material guiding hand. In this essay, I present an evolutionary pathway from so-called prokarytic cells to eukaryotic cells, and which pathway accounts for much of the complexity found in the latter. The essay follows from my reading of Duz and Dincer[1] Gabaldon[2] and Devitt.[3]

Almost all plant and animal cells contain mitochondria. Mitochondria are important little organelles because they manufacture certain highly energetic molecules which cells need to get things done. Given the importance of mitochondria, then, where did they come from? There is compelling circumstantial evidence that a deep ancestral connection exists between mitochondria and certain types of bacteria. In fact, it is most likely that mitochondria were once free-swimming bacteria. The key to this mitochondrial/bacterial connection is something called endosymbiosis.

The coming about of endosymbiosis on Earth is considered a major evolutionary transition. Some organisms evolved an ability to live inside the cells of other organisms without being digested, and such that both themselves and their hosts cells benefit. Indeed, such endosymbiotic relationships between organisms are widely observed today.

A long time ago, life on Earth was much simpler than it is today. That's because only simple organisms existed. These were the bacteria, cyanobacteria and archaea. Each would consume each other. But the cyanobacteria also evolved an ability to convert carbon dioxide into food using sunlight. Some of the bacteria which were engulfed by the archaea were not digested. Instead, they acquired an ability to coexist inside the archaeal cells. In turn, the archaeal cells became accustomed to the presence of these bacterial cells inside their cytoplasm.

Over time, both the bacteria and the archaea grew to depend on each other more and more, sometimes even exchanging fragments of their own DNA material in a process known as horizontal gene transfer. Eventually, neither the host archaea nor the engulfed bacteria could survive without the other. And so the bacterial cells eventually lost their bacterial character and evolved into mitochondria. Correspondingly, the host archaeal cells lost their archaeal character and evolved into plant and animal cells. This remarkable story is similar for how some cyanobacteria evolved to become the photosynthetic chloroplast organelles found in plants cells. This is known as endosymbiosis. Endosymbiosis thus inadvertently offered life an important evolutionary pathway for a ratcheting up of cellular complexity on Earth.

in-out-eukaryogenesis.jpg
Schematic of the inside-outside autogenous model of eukaryogenesis. In the model, an archaeon engulfs free-swimming proteobacteria by the outward expansion of its endosplasm (1, 2 & 3). The expansion begins as blebs which form at the archaeon's outer cell wall and plasma membrane (2). This outward expansion offers the proto-eukaryote cell a foraging advantage (3) because the increased surface area of its plasma membrane increases the likelihood of engulfment by phagocytosis of nearby organisms, whilst protecting its nuclear DNA material in its central nucleoid (4). Some engulfed aerobic proteobacteria resist digestion (4) and evolve into the familiar mitochondria organelles (5). Over time, the nucleoid evolves into the eurkaryotic nucleus (5 & 6). The endoplasmic reticulum, the double-layered phospholipid enveloping membrane surrounding the nucleus, and the protein complexes located at the nuclear pores are all structural and functional remnants of the process (6).

So in writing this short essay, I wish to honour those early bacteria, cyanobacteria and archaea. I thank them for their struggles to survive and thrive, and to learn how to co-exist and cooperate. But I also lament that as we inevitably dwell on our human-centric issues of today and tomorrow, we often pay little heed to the role that these and a myriad other creatures played across the aeons in making us us. Bacteria, cyanobacteria and archaea, with your endosymbiosis, please take a bow!


  1. 1 I declare this to be my own work, entirely. In particular, no AI was used in any research, analysis, synthesis, writing, nor typesetting of this work. In short, AI was not recruited at any time in this work. Errors and inaccuracies are therefore proudly my own.
  2. 2 Thanks to Leslie Viljoen whose Facebook post on 25Aug2023 inspired this essay.

  1. [1] NB Duz and P Dincer. A review on theories on the origin of the nucleus in modern eukaryotes. Gene Technology. 10(5):, 2021. Accessed 25 August 2023. https://www.walshmedicalmedia.com/open-access/a-review-on-theories-on-the-origin-of-the-nucleus-in-modern-eukaryotes-88171.html
  2. [2] T Gabaldon. Origin and early evolution of the eukaryotic cell. Annual Review of Microbiology. 75(1):, 2021. Accessed 25 August 2023. https://doi.org/10.1146/annurev-micro-090817-062213
  3. [3] Terry Devitt. New theory suggests alternate path led to rise of the eukaryotic cell. Accessed 3 September 2023. https://news.wisc.edu/new-theory-suggests-alternate-path-led-to-rise-of-the-eukaryotic-cell/

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