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Part A shows a eukaryotic cell. The illustration indicates that, within the nucleus, operational genes were inherited from an ancestral Eubacteria, and informational genes were inherited from an ancestral Archaebacteria. Part B indicates that the outer membrane of Gram-negative bacteria is derived from Archaea, and the inner membrane is derived from Gram-positive bacteria.
The theory that mitochondria and chloroplasts are endosymbiotic in origin is now widely accepted. More controversial is the proposal that (a) the eukaryotic nucleus resulted from the fusion of archaeal and bacterial genomes, and that (b) Gram-negative bacteria, which have two membranes, resulted from the fusion of Archaea and Gram-positive bacteria, each of which has a single membrane.

The nucleus-first hypothesis proposes that the nucleus evolved in prokaryotes first ( [link] a ), followed by a later fusion of the new eukaryote with bacteria that became mitochondria. The mitochondria-first hypothesis proposes that mitochondria were first established in a prokaryotic host ( [link] b ), which subsequently acquired a nucleus, by fusion or other mechanisms, to become the first eukaryotic cell. Most interestingly, the eukaryote-first hypothesis proposes that prokaryotes actually evolved from eukaryotes by losing genes and complexity ( [link] c ). All of these hypotheses are testable. Only time and more experimentation will determine which hypothesis is best supported by data.

 Part A shows the nucleus-first hypothesis. According to this hypothesis, a primary endosymbiotic event resulted in an ancestral eukaryotic cell acquiring a nucleus, and a secondary endosymbiotic event resulted in the acquisition of a mitochondrion. Part B shows the mitochondrion-first hypothesis. According to this hypothesis, the mitochondrion was acquired before the nucleus, but both were acquired by endosymbiosis. Part C shows the eukaryote-first hypothesis. According to this hypothesis, prokaryotes evolved from eukaryotic cells that lost their nuclei and organelles.
Three alternate hypotheses of eukaryotic and prokaryotic evolution are (a) the nucleus-first hypothesis, (b) the mitochondrion-first hypothesis, and (c) the eukaryote-first hypothesis.

Web and network models

The recognition of the importance of HGT, especially in the evolution of prokaryotes, has caused some to propose abandoning the classic “tree of life” model. In 1999, W. Ford Doolittle proposed a phylogenetic model that resembles a web or a network more than a tree. The hypothesis is that eukaryotes evolved not from a single prokaryotic ancestor, but from a pool of many species that were sharing genes by HGT mechanisms. As shown in [link] a , some individual prokaryotes were responsible for transferring the bacteria that caused mitochondrial development to the new eukaryotes, whereas other species transferred the bacteria that gave rise to chloroplasts. This model is often called the “ web of life    .” In an effort to save the tree analogy, some have proposed using the Ficus tree ( [link] b ) with its multiple trunks as a phylogenetic to represent a diminished evolutionary role for HGT.

 Illustration (a) shows the web of life. The base of this web is an ancestral community of primitive cells. This pool of ancestral cells gave rise to the three domains of life. However, because of gene transfer and endosymbiosis events, connections occur between the branches at various points. Thus, eukaryotic chloroplasts and mitochondria originated in bacterial lineages, and archaea and bacteria have exchanged genes.
In the (a) phylogenetic model proposed by W. Ford Doolittle, the “tree of life” arose from a community of ancestral cells, has multiple trunks, and has connections between branches where horizontal gene transfer has occurred. Visually, this concept is better represented by (b) the multi-trunked Ficus than by the single trunk of the oak similar to the tree drawn by Darwin [link] . (credit b: modification of work by "psyberartist"/Flickr)

Ring of life models

Others have proposed abandoning any tree-like model of phylogeny in favor of a ring structure, the so-called “ ring of life    ” ( [link] ); a phylogenetic model where all three domains of life evolved from a pool of primitive prokaryotes. Lake, again using the conditioned reconstruction algorithm, proposes a ring-like model in which species of all three domains—Archaea, Bacteria, and Eukarya—evolved from a single pool of gene-swapping prokaryotes. His laboratory proposes that this structure is the best fit for data from extensive DNA analyses performed in his laboratory, and that the ring model is the only one that adequately takes HGT and genomic fusion into account. However, other phylogeneticists remain highly skeptical of this model.

Illustration shows a ring with the words “pool of primitive prokaryotes” in the middle. Three arrows point outward from the ring, pointing at the three domains, Bacteria, Archaea, and Eukarya, indicating that all three domains arose from a common pool of prokaryotes.
According to the “ring of life” phylogenetic model, the three domains of life evolved from a pool of primitive prokaryotes.

In summary, the “tree of life” model proposed by Darwin must be modified to include HGT. Does this mean abandoning the tree model completely? Even Lake argues that all attempts should be made to discover some modification of the tree model to allow it to accurately fit his data, and only the inability to do so will sway people toward his ring proposal.

This doesn’t mean a tree, web, or a ring will correlate completely to an accurate description of phylogenetic relationships of life. A consequence of the new thinking about phylogenetic models is the idea that Darwin’s original conception of the phylogenetic tree is too simple, but made sense based on what was known at the time. However, the search for a more useful model moves on: each model serving as hypotheses to be tested with the possibility of developing new models. This is how science advances. These models are used as visualizations to help construct hypothetical evolutionary relationships and understand the massive amount of data being analyzed.

Section summary

The phylogenetic tree, first used by Darwin, is the classic “tree of life” model describing phylogenetic relationships among species, and the most common model used today. New ideas about HGT and genome fusion have caused some to suggest revising the model to resemble webs or rings.

Questions & Answers

Please what is ovulation
Adusei Reply
what is a metaborism
Beatrice Reply
this is a specialized part of the cell eg Nucleus
David Reply
what are the organelles?
Faith Reply
what is reproduction
mana Reply
reproduction is the process by which living organisms give rise to young ones of their own kind
Miriam
What is evolution
Wengelawit
the production of new forms of life over time as documented in the fossil record.
mana
hmm
Marvin
give two parasites where secondary host is water snail?
Kevin Reply
what is treats
Raih Reply
what are the organelles in cell that involves in protein sythenis
Rita Reply
what is a melanin?
Judith Reply
what is telophase
Elphas
melanin in that black color we posse in our skins
Marvin
Why do parasites take on a parasitic life?
Asadullah Reply
what is cell
Ousman Reply
A cell is the basic structure unit of an organ
Yaa
what is respiration
Igwe
what is organisms?
Bashir
Organisms are living things that can function on their own
EZEA
A cell is a functional basic unit of an organisms.
Evelyn
Organisms is a entity which consists of one or more cells and are able to undergo all life processes
Evelyn
A respiration is the physical and chemical break down of complex food substance into absorbable or simple form.
Evelyn
What will to a cell if the nucleus is removed..?
Goodrick Reply
When a cell nucleus removed then the cell will not be able to function properly, it will not be able to grow .All the metabolic functioning of the cell will stop .Without nucleus the cell will lose it's control. It can not carry out cellular reproduction .
Evelyn
defin peroxesome
walid
write a short note on how the kidney carry out osmoregulation in man
Rhoda Reply
Kidney play a very large role in human osoregulation by regulating the amount of water reabsorbed from glomerular filtrate in kidney as tubules, which is controlled by hormones such as antrdiuvetic hormone (ADH) , aldosterone and angiotensin.
Evelyn
kidney is capable of osmoregulation in mammal ,
Judith
simple definition for respiratory system
Gift Reply
Respiratory system is a network of organs and tissues that helps you to breathe or help in getting the rids of oxygen and discharge of carbon dioxide in the body.
Ruhiyatu
What are uses of respiratory system
Joster
how it functions
Ousman
what is inresparetion
Fikkabex
Why do parasites take on a parasitic life?
Asadullah
A respiratory system is a biological system consisting of specific organs and structures used for gas exchange in animals and plant
Evelyn
diseases of respiration
walid
when air enters to the body called inresparetion
walid
explain why plants responds to stimuli slowly than animals
Bekoe
how is a aerenchyma tissue adapted to its function
fatuma Reply
Have large air spaces that store air for gaseous exchange... Large air spaces also facilitate bouyancy.
Goodrick

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Source:  OpenStax, Biology. OpenStax CNX. Feb 29, 2016 Download for free at http://cnx.org/content/col11448/1.10
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