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Bio 150 Molecular Cell Biology - Multicellular prokaryotes not mentioned in the figure

Miriam de Vera, PhD In the figure, prokaryotes may be single unicellular


Lecture 1.3 Cell Evolution (Evolution of the beginnings of life) organisms or colonial

Earth 3. Proterozoic Eon


- Approximate age of the earth: 4.5 BYO - Oxygenated atmosphere needed prior to the evolution of
- Conditions when it was formed: Eukaryotes
Extreme - Before you can have the major phyla of organisms, that
Many volcanic activities (pyroclastic) would have to be past 1 billion years ago
Rough waters - Aquatic before terrestrial life forms
Thunderstorms - Plantae most recent division to emerge
Extremes of temperature (fire) - 2.5 billion years atmospheric O2
Crust of earth almost molten so we dont expect that Animals first formed before vascular plants
exactly 4.5 BY ago, theres already life Colonization of land insects
A lot of geologic activities going on - 1.75 billion years single celled eukaryotes
- 1.5 billion years multicellular eukaryotes
Major episodes in the history of life
Evidences for these claims
- Earliest cells are most likely prokaryotic
All life begins from a simple cell prokaryotic
- Stromatolites (figure)
Banded domes or mats of sedimentary rocks
Age: roughly 3.5-3.8 BYO suggesting the earliest time
that prokaryotes evolved
Oldest known stromatolites: 3.8 BYA; rock materials
which contain evidence of ancient prokaryotes
Majority can be found in marine environments or near
coastal areas
Cross sections: layers/bands can be seen which are
typical of sedimentary rocks; these bands are from the
layering of the biofilm of the prokaryotes
Thrombolites: subtype of stromatolites; specifically
lain down by photosynthetic microorganisms. These
are not necesarilly photosynthetic
*no need to memorize geologic timescale Figure: Reddish bands layers of iron oxides
suggesting that the process of oxidation is already
1. Origin of solar system occurring to promote the layering some traces of
- 4.5 BY ago O2 are being generated by some of these bacteria
- Theres a mass at the very center which became the sun (although not yet fully integrated into the atmosphere
- Surrounding the sun are coalescing particles which since they are still on the rock layers)
comprise the planets
- Gravitational pull for the circling of planets is provided by the
central source of energy SUN
- Accretion Theory
One theory for the formation of solar system
Basin of water: represents space
Pour talcum powder on the surface only (do not
disturb water)
At the very center, stir using a stick (represents the
Minerals formed from energy processes of cells
rotation of the sun)
Columnar, deeper in the oceans
A mass will form at the center while smaller masses
Orange rocks: calcium and magnesium
at the periphery will occur
Certainly, many of those particles obtain solid mass
Supp. Notes: Carbon isotopes (representing metabolic activity of
unlike the peripheral planets, they are mainly gaseous
organisms) have been found in 3.5 BYO rocks.
2. Archaean Eon
Something should have been present between formation of earth
- Prokaryotes 3.8 BYO (earliest organisms), oldest life
and existence of cells/prokaryotes (4.5 BYA 3.8 BYA)
forms on earth; based on radiocarbon dating of certain
- Many molecules formed (not only inorganic rocks)
rocks that are 3.8 BYO
- Simple molecules more complex (biomolecules)
- Earliest prokaryotes did not utilize oxygen

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Chemical Evolution - Miller and Urey [1953]
- The first cells may have arisen by chemical evolution on a Attempted to simulate primitive earth conditions to
young Earth spontaneously produce organic cpds from
- inorganic + organic compounds earliest life forms inorganic substances
- Inorganic organic molecules/compounds Initial mixture containing inorganic substances
- Stages: (water, liquid N, hydrogen, oxygen, etc. no
1. Abiotic synthesis and accumulation of organic hydrocarbons at all) subjected to intense heat
molecules [monomers] simulating intense kinetic energy in primordial
o Should be MONOMERS first earth exposed to electrical energy
o Requires high energy reactions from the environment After letting the set-up ran for a while, they
synthesis of organic molecules collected pure liquid from the apparatus
2. Assemblage of monomers into polymers chemical analysis hydrocarbons were found
3. Aggregation of abiotically produced monomers or NOTE: no cells, enzymes, etc were used, only
polymers into protobionts (protocells) kinetic energy and inorganic cpds
o Protobionts - aggregates of abiotically produced Figure: CH4 (methane) simplest organic
molecules surrounded by a membrane or molecule
membrane-like structure; not alive yet; seemingly Before the sample for chemical analysis, there
resemble cells were already some preformative polymers
o Experiments showed that protobionts can be There were others who replicated the expt and they
manufactured synthetically produced different organic cpds (simple
4. Origin of heredity monomers/oligomers)
o Heredity transmission of information
o Note: NON-LIVING

Prebiotic Chemical Evolution


1. Abiotic synthesis and accumulation of organic molecules
[monomers]
- Organic
they just simply form from strings of carbon
attached to hydrogen hydrocarbons (CH)
inorganic in origin
- Before this, only inorganic molecules are present which
are essentially elements by themselves or small
molecules without carbon
- C, H, O (little), N and other low mass elements
- Oparin and Haldane [1920s]
conditions on primitive earth favored chemical
reactions that synthesized organic compounds
from inorganic substances 2. Assemblage of monomers into polymers
Conditions: extreme, highly kinetic envt. (lighting, - Covalent bonds, hydrogen bonds
hurricane, volcanic activities, etc.) wherein - Could not possibly spontaneously happen (no
inorganic molecules in any envt will tend to clash involvement of any biotic elements)
with one another - Ancient conditions still apparent, particularly heat
1 billion years of clashing together in random will - Laboratory simulations of early earth conditions have
ultimately form hydrocarbons linked together produced organic polymers
ORGANIC compounds - S. Fox [U. Miami] produced protenoids (polymers)
There are chemical reactions everywhere that from dilute solutions of organic monomers (amino
favor these collisions of molecules to form organic acids) applied unto hot substrate
compounds (monomeric). Most likely, they will be Protenoids not cellular in origin; not fully
stored up in the sea (water bodies) translated by cells unlike proteins
PRIMORDIAL SOUP (where the 1st monomeric Polymers formed through exposure to heat
organic compounds may have formed) - G. Wachterhauser certain properties of pyrite (fools
- Oxidizing atmosphere of present earth not conducive gold) have catalyzed abiotic synthesis of organic
to spontaneous synthesis of complex molecules due to molecules
electron extracting nature of O2 (destabilizing chemical Pyrite heat absorbent and with electrical
bonds) properties; if papatak-patakan ng monomers
much more diversity of organic compounds such as amino acids, polypeptides can be formed
spontaneously on the surface over time
Note: in very extreme conditions (as in SUPER
HOT!!!)

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- These 2 scientists mentioned showed that proteins can A. Liposomes
be formed abiotically - Very similar characteristic to a soap bubbles
- Have simple reproduction in a sense that it can be
3. Aggregation of abiotically produced monomers into amplified (like blowing soap bubbles, press mo lang
protobionts magiging marami na)
- Types of Protobionts (complex of molecules inside a - The fact that they can reproduce (chem nature), shows
membrane): that they somehow exhibit some living properties
A. Coacervates
combination of B. Coacervate
carbohydrates and - can be manufactured in the lab (using glucose
proteins or phosphate)
carbohydrates and - enzymes isolated
lipids; can have 2 - no need for triggers to activate enzymes, it can be injected
layers of membrane there or it might have been produced from some external
protein translation process (basta makapasok lang, after
B. Microspheres all, membrane is semipermeable)
proteins or amino - starting material + enzymes chemical process occurs
acids bounded by a - chemical reactions reflective of what happens in actual
membrane; used for living cells properties of living cells simulated
transport; used to - Coined by Oparin
introduce proteins to - Colloidal organic compounds (usually fatty acids) bounded
tissue and cells by membrane

C. Liposomes contains lipids (mostly 4. Origin of heredity


phospholipids) or fatty acids; semi-permeable
membrane properties; can transfer contents from
one liposome to another
- In certain industrial companies, they produce
microspheres and liposomes to serve as carriers of
proteins or fatty acids which are transferred to another
chemical batch (in manufacturing process esp in
pharmaceuticals)
- Protobionts may look cell-like but they are not alive;
exhibit some of the properties associated with life,
including:
Simple reproduction
Metabolism
Enzymes introduced into liposomes
enzymes released - Heredity - some molecular information must be passed
With right conditions (though non-living), can on
undergo processes - There should be some means of a molecule being
Maintenance of an internal chemical envt. reformed
different from that of their surroundings - Among all of the various kinds of polymers
- Earliest ones evolved spontaneously (biomolecules), it is believed that the nucleic acid (RNA
not DNA) is the best kind of molecule that can transmit
Properties of living cells can be simulated using protobionts information
- RNA served as the first hereditary molecule, subject to
spontaneous reassembly
DNA: more complex and stable than RNA;
requires more components in order to be formed
- RNA can replicate themselves (self-replication)
DNA: still needs DNA polymerase, helicase, etc.
- Figure: Earliest kind of RNA replication
- Pink and blue: RNA
- Pink: single stranded but with bends (loops) making
some parts double stranded; RNA molecule functioning
as a ribozyme
- Ribozyme = enzyme & template
enzyme for producing its own copy and
template for the assemblage of nucleotides

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- Becomes a copy of a copy
- Ribozyme only. No need for other complex
accompaniments for the formation of complementary
RNA copy
- There are different kinds of ribozymes which can be used
for RNA replication
- Following the simplicity-complexity rule, RNA (simple) is
logically the first to arise before the DNA (complex)
- Even short pieces can be used for heredity

Picture: Sampling the water around a hydrothermal vent in the Sea


Supp Notes: Thomas Cech, of the University of Colorado, and
of Cartes. More than 1.5 km below the surface, the vent releases
Sidney Altman, of Yale University, found that RNA, which plays a
hydrogen sulfide and iron sulfide, which react and produce pyrite
central role in protein synthesis, can also carry out a number of
and hydrogen gas. Prokaryotes that live near the vent use the
enzyme-like catalytic functions. Cech called these RNA catalysts
hydrogen as an energy source. Such environments are among the
ribozymes. Some ribozymes can make complementary copies of
most extreme in which life exists today, and some researchers
short pieces of RNA, provided that they are supplied with
favor the hypothesis that life may have begun in similar regions of
nucleotide building blocks.
early Earth.

Present Trends (Transition from non-living to living)


Simpler self-replicating molecules (other than RNA)
Panspermia and Chemical Evolution [17th C.]
Trebek et al. [1991]
- Suggests that the earliest life forms might have had an
- produced aminoadenosine triacid ester (AATE)
extraterrestrial source
Has a composition and properties for self-replication
Life on earth came from somewhere else in the
(essential characteristic of heredity)
universe
Very simple, rel. small molecule
- Meteorites
- Figure: amino adenosine and ester can be combined through
Extraterrestrial
heat or any anabolic process to form a template molecule
If proven that there is no contamination on these
- If you have a mixture of several of these molecules, one can
meteorites (from organic materials found on earth),
serve as template (see fig)
they may have had organic materials coming from
- Rightmost fig: when condition is reversed, ex. Cooling
outside of the Earth
you can have replicas (exactly the same)
Naturally, only inorganic compounds are found in
- No enzymes
meteorites. Surprisingly, organic cpds are found in
- Caused by extreme agitation
meteorites.
- So, there might be other molecules smaller than RNA that
Probably these cpds brought about the first
could have this property provided that you can provide the
prokaryotes from protobionts
necessary conditions for their formation (abiotic)
- It is possible that at least some organic compounds reached
the earth from space in the form of meteorites or comets
hitting the earth (extraterrestrial origin)

Wachterhauser et al. [1970s]


- Explored hydrothermal vents (very deep water)
- Hydrothermal vents
Represent some of the most extreme environments on
earth F. Dyson
Mirror the extreme conditions of the earths marine envt - Has suggested that the 1st RNA molecules may have been
in the past short, virus-like sequences (aided in their replication by
- Samples of organic polymers and prokaryotes are gathered random amino acid polymers that had rudimentary catalytic
from these hydrothermal vents capabilities)
- No protobionts found yet (so theres still no transitional - short virus-like sequences might be the earliest self-
evidence between molecules and cells) replicating molecules (if no other molecules are self-
- Its possible that the process of evolving is still happening so replicating)
we should continue to explore these hydrothermal vents
- Organisms from hydrothermal vents Earliest Prokaryotes
H2S + Fe2S Pyrite + H2(g) - Originated 3.5-4 billion years ago
- Newer species of archaea bacteria from extreme environments - Metabolically simple with few enzymes
- Most likely: anaerobic and opportunistic
Anaerobic: no O2 when they first evolved
Opportunistic: utilized whatever resources is
abundant; usually facultative

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Mode of Nutrition (theyre alive and can metabolize already! ) Supp Notes:
a) An early chemoheterotroph lives on ATP and other
organic compounds absorbed from its surroundings.
Enyzmatic breakdown of the organic compounds
Inorganic provided energy and carbon.
b) Earliest mode of energy metabolism was
Inorganic chemoautotrophic (among autotrophs). Ancestral cells
obtain carbon from CO2 and synthesize their own ATP by
simple chemiosmotic mechanism. The model shows
enzymes promoting chemical reactions among simple
inorganic cpds. Hydrogenase couples the release of
energy from inorganic compounds to the splitting of H2
and maintenance of a proton gradient across the plasma
membrane. The potential energy of the proton gradient
drives the enzymatic synthesis of ATP.
*Heterotrophs cant produce organic cpds from inorganic cpds
**Eventually, certain prokaryotes have evolved systems which
utilize light Contain photosynthetic pigments

Photosynthetic Pigments found among prokaryotes


- Examples (figure):
Bacteriorhodopsin found in purple sulfur bacteria
(use H2S rather than H2O)
Chlorophyll

Heterotroph

Bacteriorhodopsin Chlorophyll
- Light absorbing pigments may have been coupled with
membrane proteins involved in ATP synthesis
The light that is utilized in these processes for the
Autotroph photoautotrophs drive the machinery for proton
pumps (for ATP synthesis)
- Heterotroph mode of nutrition which evolved first among - Photosynthesizing bacteria evolved later after the anaerobic
prokaryotes chemoheterotrophs
ATP organic - Photosystems may have evolved that use light to drive
From simple breakdown, you can immediately have electrons from H2S to NADP+, generating the reducing power
energy to fix CO2
Organic cpds polymers in the envt; present even H2S not H2O since theres still no oxygen (later,
before protobionts existed; used by heterotrophs organisms which utilize water will arise)
- Autotroph NADP+ nicotinamide adenosine dinucleotide
More complex phosphate; oxidized form of NADPH
Needs transporter, proton gradient, means of CO2 fixation formation of carbon-based compounds
utilizing energy to fix CO2 to form hydrocarbon - Prokaryotes do not have internal membranes yet so these
The process of producing organic material from proteins and machinery are associated with the inner side of
inorganic sources is a more complex process so it the plasma membrane (no mitonchondria and chloroplast)
evolved later than heterotrophs Kailangan may kapitan yung machinery for ETC so
Opportunistic: can use inorganic sources these proteins are attached/associated with the PM
- The earliest prokaryotic ancestors are likely to be - Membrane proteins that powered ATP synthesis may have
anaerobic heterotrophic cells been opted
- Their energy has to be derived from food materials (pre- Co-option change of a structure with an initially
made source of energy from outside their body. different function
- They can exist without molecular oxygen

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Supp Notes: - Evolution of anti-oxidation systems
Bacteriorhodopsin absorbs light and uses the energy for H+ O2 attacks the bonds of organic molecules
pumping to drive ATP synthesis (simplest mechanism for (destructive)
photophosphorylation). Membrane proteins that powered atp Antioxidant systems evolved to tolerate rising O2
synthesis may have been co-opted to provide the reducing power levels
to fix CO2 (e.g. bateriochlorophyll). Proteins that can sequester away oxygen to prevent it
from inhibiting some metabolic processes
Purple sulfur and green sulfur bacteria split H2S instead of H2O. - Modification of ETC co-opted from photosynthesis (resulting
in anaerobic respiration)
First Cyanobacteria Aerobic respirations may have originated as a
modification of ETC co-opted from photosynthesis.
O2 - final e- acceptor in ETC
Co-opted - from the earliest cyanobacteria, it gets
modified to become functional even in the obligate
anaerobes
- Resorting to chemoheterotrophic nutrition most recently
- Figure: evolved
Left Eutrophic lake (bluish greenish because of the Chemoheterotrophs adapt the ETC to function
presence of cyanobacteria) exclusively in aerobic respiration. (e.g. purple sulfur
Right Anabaena; filamentous colonial cyanobacteria bacteria)
Note: earliest prok are non-colonial or solitary
- Had the metabolic machinery to use H2O instead of H2S or Natural selection tend to favor increasing complexity of earliest
other compounds as a source of electrons and hydrogen for cells
reducing CO2 for CO2 fixation to make sugars, - Single, unicellular Multicellular prokaryotes
carbohydrates inside the cells Natural selection favor colonial species
- Anabaena Colonies (fig) Advantage: specialization; division of labor
Multicellular Differentiation of function (more functions = better
Some photosynthetic, involved in CO2 fixation natural selection will favor them)
similar cells with pigments (near the arrowhead) - Complex bacterial communities
but there are others that are not heterocytes Aggregate of different kinds of species co-existing
- Heterocytes together
multicellular/colonial Biofilms some of the bacteria here can produce
cells that are different from other cells in the Anabaena signaling molecules for other members of the
colony community to gather or stay away; material containing
involved in nitrogen fixation, not photosynthesis various bacterial populations and exudates [chemical
large; have thick cell walls to prevent diffusion of O2 signals]
because O2 is inhibitory to nitrogen fixation which is Leads to more diversity among prokaryotes
necessary for the manufacture of amino acids and - Compartmentalization of functions within a cell
nucleotides Prokaryotes do not have internal organelles at this
- Implication of colonial/multicellular arrangement point yet
can have specialization of function Localization of proteins in the PM
some are photosynthetic, some are involved in N2 Compartmentalization is implied by the position of
fixation certain proteins in some specific points of attachment
they can live in envt with minimal resources (just like in the PM
those in eutrophic lake) Proteins for ETC transport, protein sorting, and
other metabolic transport assoc with the PM
**green fur of mammals in moist/wet areas due to blue-green Complete compartmentalization involves the
algae growing on their fur evolution of separate compartments organelles
**Nostocaceae family of Anabaena Hence, this would necessarily be the step leading to
**Atmosphere became more oxidizing than reducing due to the evolution of eukaryotes
increasing photosynthetic organisms (high O2)
Eukaryotic Origins
Consequences of O2 in the Atmosphere for Prokaryotes Acritarchs
- Extinction of anaerobes - earliest fossils of eukaryotes
Facultative anaerobes can survive; prefer anaerobic - oldest one found from the fossil record is approx. 2 BYO
conditions but can thrive in the presence of O2 - resemble cysts of protists (spherical; thick capsule)
- Evolution of obligate anaerobes - Earliest protists [eukaryotes from gradual evolution of more
There are still environments which are anaerobic. They complex structures]
can evolve here
Bacteria are ubiquitous

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Mechanism of the development of internal membranes in - Ancestral heterotrophic prokaryotes (early beginnings)
eukaryotic cells: have a machinery similar to the ATP producing
machinery of the mitochondria
Heterotroph hid inside the larger ancestral cell for
protective measure. They just remained inside
because they benefit each other naman
Heterotroph can get protection; Host cell can get
energy
Large ancestral prokaryote engulfed heterotrophic
- Plasma membranes extent of infolding internally becomes prokaryote harnessed ATP of prokaryote early
more and more complex organelle: ER eukaryote with mitochondria
- Hypothesis: ER contributes to the formation of porous Heterotrophic eukaryotes Animalia
nuclear membrane compartmentalization for the genome
- These internal membranes that are formed are part of the - Ancestral photosynthetic prokaryotes
endomembrane system so theres still no exact discrete precursor of chloroplasts
compartments (continuous pa rin because of pores) Large ancestral prokaryote engulfed photosynthetic
- The endomembrane system of eukaryotes including the nuclear prokaryote eukaryote with chloroplast
membrane may have evolved by the in-folding of the plasma
membrane of an ancestral prokaryote.
1800s: Russian scientist Mereschkowski presented the
- Diagram:
endosymbiont theory
Membrane-enclosed compartments
1970s: Lynn Margulis revived the idea. She provided the molecular
PM develops in-foldings branching
and genetic evidences.
Compartments detached from PM
Supp Notes:
Supp. Notes: The endomembrane system of eukaryotes including Mitochonrdria and chloroplasts evolved from prokaryotes that lived
the nuclear membrane may have evolved by the infolding of the symbiotically within some ancestral eukaryotes. The similarities in
plasma membrane of an ancestral prokaryote. appearance between these structures are pretty clear. The inner
membrane of the mitochondrion protrudes into the interior rion in
EndosymbioticTheory
folds called cristae. This membrane is very similar in appearance
- Hypothesis that can exemplify the formation of the more
to the prokaryotic plasma membrane with its mesosomes.
discrete organelles for the eukaryotic lineage
Origin of Eukaryotes
[Refer to the article Origin of Eukaryotes (from Maam DV)]
See discussion on engulfed heterotrophic and photosynthetic
prokaryote

- Mesosomes found in prokaryotes; similar to the infoldings


in the mitochondria (cristae)
- Cristae more complex branching of infoldings
- Figure:
Ancestral host cell with endomembrane system and
apparent nucleus
More discrete organelles
- Margulis modern proponent of endosymbiont theory;
revived the idea of the concept that:
the cause of having discrete organelles in eukaryotic
cells is due to the entry of symbiotic bacteria into the
cytoplasm of the larger host cell

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Figure: A phylogenetic tree based on rRNA sequence comparisons
showing the three domains of life. The Archaea are divided into two **PNAS Proceedings of the National Academy
subgroups as indicated. - 13 organisms
- Similarity index (SI) used to compare extent of similarity
Tree of Life The higher the value, the greater the similarity in the
- 3 domains: Archaea, Eubacteria, Eukarya sequences
- Previously K. Monera has been elevated to 2 separate - 1 to 3: Eukarya (yeasts, animal cell line)
domains: Archaea and Eubacteria - 4 to 9: Eubacteria (bacteria, chloroplast with own RNA)
- As seen in the grouping, Archaea and Eukarya are more closely - 10 to 13: Archaea (extremophiles)
related - From the 13 organisms that were used as test subjects, they
They have a common ancestor were essentially divided into 3 distinct groups
- Archaea includes extremophiles, the most ancient of - However, there are other studies which showed more similar
prokaryotes nucleotide sequences between Archaea and Eukarya than
- Nucleotide sequences - molecular evidence used for the Euk and Eubac or Archaea and Eubac
relatedness of Eukarya to Archaeabacteria - Informational molecules used: nucleic acids and proteins
used in transcription, translation, replication
- Informational molecules more reliable than metabolic ones
Metabolic proteins are not used (ex. cytochrome
proteins)
Lateral gene transfer
Ex. exchanging genes thru pilus
they do not transfer genetic info by
progeny/reproducing like cell division or cell
fission
Instead, they happen to pick up certain genes
from different species in which they share a
common environment
Therefore, it is possible that some genes from
the Archaea could have been mixed with
Eubacteria or vice versa (As well as Eubacteria
and Eukarya)
Usually such genes or molecules are metabolic
- 2D electrophoretic gel of the components of small subunit in nature
ribosomal RNA from 13 different species (see next table) Vertical gene transfer from parents to offspring
- 2 dimensional bc 2 step separation: (progeny); informational molecules
1st run: pH 3.5; based on size
2nd run: pH 2.3; based on charges Di sure si maam ditto. Just read the article nalang.
- Separated extracted determine nucleotide sequence for - By comparing the nucleotide sequence of small subunit
the different species rRNA, it shows that there are 3 very distinct evolutionary lines
- Woese and colleagues showed evidences on which - The table suggests that beyond the domains, the values with
organisms are more related to one another the next succeeding highest levels of SI are shared between
Depends on shared molecular characteristics Eukarya and Archaea
More similar = more related
molecules involved are informational molecules like
RNA and proteins; not metabolic molecules like carbs
and lipids
- READ THE ARTICLE

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Comparing the Three Domains

- 3 domains occasionally have some similarities


- The greatest genetic similarities (sequences of info mol) are
between Archaea and Eukarya even though eukaryotic line
evolved later after Eubacteria line

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