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Evolution of Cells PDF
Evolution of Cells PDF
Cells are divided into two main classes, initially defined by whether they contain a
nucleus. Prokaryotic cells (bacteria) lack a nuclear envelope; eukaryotic cells have
a nucleus in which the genetic material is separated from the cytoplasm.
Prokaryotic cells are generally smaller and simpler than eukaryotic cells; in
addition to the absence of a nucleus, their genomes are less complex and they do
not contain cytoplasmic organelles or a cytoskeleton . In spite of these
differences, the same basic molecular mechanisms govern the lives of both
prokaryotes and eukaryotes, indicating that all present-day cells are descended
from a single primordial ancestor. How did this first cell develop? And how did the
complexity and diversity exhibited by present-day cells evolve?
Prokaryotic
The First Cell
It appears that life first emerged at least 3.8 billion years ago, approximately 750
million years after Earth was formed.. How life originated and how the first cell
came into being are matters of speculation, since these events cannot be
reproduced in the laboratory. Nonetheless, several types of experiments provide
important evidence bearing on some steps of the process.
Time scale of evolution.
Time scale of evolution. The scale indicates the approximate times at which some
of the major events in the evolution of cells are thought to have occurred.
It was first suggested in the 1920s that simple organic molecules could form and
spontaneously polymerize into macromolecules under the conditions thought to
exist in primitive Earth's atmosphere. At the time life arose, the atmosphere of
Earth is thought to have contained little or no free oxygen, instead consisting
principally of CO2 and N2 in addition to smaller amounts of gases such as H2, H2S,
and CO. Such an atmosphere provides reducing conditions in which organic
molecules, given a source of energy such as sunlight or electrical discharge, can
form spontaneously. The spontaneous formation of organic molecules was first
demonstrated experimentally in the 1950s, when Stanley Miller (then a graduate
student) showed that the discharge of electric sparks into a mixture of H2, CH4,
and NH3, in the presence of water, led to the formation of a variety of organic
molecules, including several amino acids (Figure 1.2). Although Miller's
experiments did not precisely reproduce the conditions of primitive Earth, they
clearly demonstrated the plausibility of the spontaneous synthesis of organic
molecules, providing the basic materials from which the first living organisms
arose.
Self-replication of RNA.
Self-replication of RNA. Complementary pairing between nucleotides (adenine [A]
with uracil [U] and guanine [G] with cytosine [C]) allows one strand of RNA to
serve as a template for the synthesis of a new strand with the complementary
sequence.
The first cell is presumed to have arisen by the enclosure of self-replicating RNA in
a membrane composed of phospholipids (Figure 1.4). As discussed in detail in the
next chapter, phospholipids are the basic components of all present-day
biological membranes, including the plasma membranes of both prokaryotic and
eukaryotic cells. The key characteristic of the phospholipids that form membranes
is that they are amphipathic molecules, meaning that one portion of the molecule
is soluble in water and another portion is not. Phospholipids have long, water-
insoluble (hydrophobic) hydrocarbon chains joined to water-soluble (hydrophilic)
head groups that contain phosphate. When placed in water, phospholipids
spontaneously aggregate into a bilayer with their phosphate-containing head
groups on the outside in contact with water and their hydrocarbon tails in the
interior in contact with each other. Such a phospholipid bilayer forms a stable
barrier between two aqueous compartments—for example, separating the
interior of the cell from its external environment.
Enclosure of self-replicating RNA in a phospholipid membrane. The first cell is
thought to have arisen by the enclosure of self-replicating RNA and associated
molecules in a membrane composed of phospholipids. Each phospholipid
molecule has two long hydrophobic.
The enclosure of self-replicating RNA and associated molecules in a phospholipid
membrane would thus have maintained them as a unit, capable of self-
reproduction and further evolution. RNA-directed protein synthesis may already
have evolved by this time, in which case the first cell would have consisted of self-
replicating RNA and its encoded proteins.
The Evolution of Metabolism
Because cells originated in a sea of organic molecules, they were able to obtain
food and energy directly from their environment. But such a situation is self-
limiting, so cells needed to evolve their own mechanisms for generating energy
and synthesizing the molecules necessary for their replication. The generation
and controlled utilization of metabolic energy is central to all celactivities, and the
principal pathways of energy metabolism are highly conserved in present-day
cells. All cells use adenosine 5′-triphosphate (ATP) as their source of metabolic
energy to drive the synthesis of cell constituents and carry out other energy-
requiring activities, such as movement (e.g., muscle contraction). The
mechanisms used by cells for the generation of ATP are thought to have evolved
in three stages, corresponding to the evolution of glycolysis, photosynthesis, and
oxidative metabolism . The development of these metabolic pathways changed
Earth's atmosphere, thereby altering the course of further evolution.
Lamarckism
Lamarckism, a theory of evolution based on the principle that physical changes in
organisms during their lifetime—such as greater development of an organ or a
part through increased use—could be transmitted to their offspring. The doctrine,
proposed by the French naturalist Jean-Baptiste Lamarck in 1809, influenced
evolutionary thought through most of the 19th century. Lamarckism was
discredited by most geneticists after the 1930s, but certain of its ideas continued
to be held in the Soviet Union into the mid-20th century.
Acquired Characteristics
Biologists define an acquired characteristic as one that has developed in the
course of the life of an individual in the somatic or body cells, usually as a direct
response to some external change in the environment or through the use or
disuse of a part. The inheritance of such a characteristic means its reappearance
in one or more individuals in the next or in succeeding generations. An example
would be found in the supposed inheritance of a change brought about by the use
and disuse of a special organ. The blacksmith’s arm (or any other set of muscles)
enlarges when used continually against an external resistance, such as the weight
of the hammer. If the effect were inherited, the smith’s children at birth would
have unusually large arms—if not at birth, then when they became adults, even
though they had not used their arms excessively. There is no evidence supporting
this case. A more subtle illustration is found in the supposed inheritance of an
increased dexterity of the hands of a musician through practice. The skill
acquired, although causing no visible increase in the size of the fingers, might be
imagined to be passed along to the musician’s children, and they might then be
expected to play skillfully with minimal practice. Just how the intricate interplay
of cerebral sequences that has given the dexterity to the musician’s fingers could
ever be transferred to the musician’s sex cells (spermatozoa or ova), and through
them to any potential children, has never been brought within the range of
biological possibilities.
Lamarckism giraffe
Jean-Baptiste Lamarck proposed that acquired characteristics were inheritable.
For example, as a giraffe stretches its neck to browse higher in trees, the
continuation of the habit over an extended period results in a gradual lengthening
of the limbs and neck.
The examples that Lamarck gives to illustrate his doctrine are illuminating. In
animals, as stated above, a new environment calls forth new needs, and the
animal seeks to satisfy them by making some effort. Thus, new needs engender
new habits, which modify the parts. The effects are inherited. For example, the
giraffe, seeking to browse higher and higher on the leaves of trees on which it
feeds, stretches its neck. As a result of this habit, continued for a long time in all
the individuals of the species, the giraffe’s front limbs and neck have gradually
grown longer. Birds that need to rest on the water—i.e., to find their food—
spread out their feet when they wish to swim. The skin becomes accustomed to
being stretched and forms the web between the toes. The horns of ruminants
have resulted from the ruminants’ butting their heads together during combats.
These examples, which appear naive in light of later discoveries, constitute some
of the evidence on which Lamarck rested his theory.
Natural Selection
Natural selection is the term that’s used to refer to the natural evolution over
time of a species in which only the genes that help it adapt and survive are
present. This idea was reported by Charles Darwin, the researcher behind many of
our modern concepts of evolution. In natural selection, a population will show
genetic traits over many generations that help it remain best suited to its
environment. These can be physical, structural traits like a skeleton or
musculature that helps it live in that setting, or can even by physiological traits
such as the presence of an enzyme in the digestive tract to help it break down the
available food sources.
Examples of natural selection
1. Skeletal Adaptations
Giraffes, lizards, and many other known species adapted to their environments
through genetic changes to their skeletons. This form of natural selection meant
that members of the population who didn’t develop and present these skeletal
changes died out. For example, giraffes developed long necks to reach food
sources higher up in trees, so members of the giraffe population who didn’t
develop a long neck died out. At the same time, certain lizards in one region
developed longer leg bones to help it climb up during periods of flood and to
escape predators in the ground; shorter legged lizards of the same population
died out until only the lizards with the long legs survived..
2. Coloration
Many species have been studied who’ve adapted to their environment through
adaptions in coloring. Once the optimal coloration is present, natural selection
occurs when members of the species without the adaptive coloring died out more
quickly and therefore didn’t reproduce as abundantly. Some example include the
deer mouse, the peppered moth, and the peacock.
3. Bacteria
Bacteria are a common research subject when studying evolution and adaptation
because some colonies of bacteria can produce several generations in one day,
letting researchers see a “fast forward” version of evolution and natural selection.
Some observed bacteria have included some who’ve adapted to new food sources
that were previously unusable, as bacteria that have adapted to the presence of
deadly antibiotics and exhibited traits that let them not only survive, but
reproduce to generate offspring that are also resistant to the antibiotic.
4. Physiological
Different species go through changes over time that help them adapt to different
environments, and humans are no different. One of the physiological changes
that different groups of human beings have made involves the ability to digest
cow’s milk. In regions where cattle are not raised, the human population is often
lactose intolerant, lacking the enzyme to break down the milk. However, in
regions where cattle are grown domestically and their milk is used as a chief part
of the food supply, those humans as a whole produce the enzyme needed to
digest milk.
Emergence of eukaryotes
Theory of symbiogenesis
The concept of symbiogenesis was introduced in 1909 by the Russian biologist
Constantin Merezhkowsky as “the origin of organisms by the combination or by
the association of two or several beings which enter into symbiosis”. In this article
we develop this idea, associated to the Freeman Dyson’s hypothesis, applied to
the early evolutive stages of life, considering that it could be a possible main rule
in the appearance and development of life conditions on Earth and elsewhere. A
cooperative, synergistic strategy should be considered as having been the
determinant in the development of the survival of the fittest, especially under
extremely adverse environmental conditions. This concept must be also applied
to the first communities of cells as the base supporting evolution of the early
“tree of life”. Cells, like we have previously described, can be included in a new
cellular concept entitled, “symbiocell”, since survival of the community under
such adverse conditions required a cooperative, synergistic strategy. Similar
principles could also be used to understand chemical pre-biotic evolution. We
believe that astrobiologists should consider it as a new approach to understand
organic and biological evolution.
1. INTRODUCTION
In his classic 1941 paper “Astrobiology”1, Laurence J. Lafleur discusses the
conditions of the origin of life in the universe, pointing out that “the process of
evolution depends upon de gradual accretion of relative minute variations or
mutations, and the occurance of these evolutionary elements is recognized to be
a matter of chance”. The author also discusses the problem in a broader
multitude approach than the quoted phrase, concluding “that life in the universe
is not confined to our planet”. Even if it was written in the 40’s, this perspective,
related to the problem of origin of life in the universe, represents an exception to
the major scientific thought of that time and the rest of the century. It also
includes, however, some limitations to the subject, not only in the scientific and
technological knowledge, as expected, but also in the epistemological way we
must understand the evolution of life. Based on the neo-Darwinian theory that is
the main approach we can offer when we look at the origin of life. Still, we believe
that other approaches could be implemented, namely using symbiogenic
principles. It is on this new vision that this paper will focus.
Both these two theories are built on the belief that evolution was a Darwinian
strict process without any cooperative or synergistic approach. However, we
believe that this approach must evolve in a more symbiogenic way, introducing
new concepts that enable us to understand the natural world in a more
cooperative sense and eventually closer to reality. In this sense, the conventional
“survival of the fittest” cannot be applied in that traditional way. A cooperative,
synergistic strategy should be considered as having been the determinant in the
development of the survival of the fittest, especially under extremely adverse
environmental conditions. One good example is the stromatolites, considered as
the first ecological strategy evolved for survival on primitive Earth, not for one
organism but for a community of organisms. This view can also be applied to the
pre-biotic evolution. In 1983, Freeman Dyson presented a hypothesis for the
origin of life using symbiogenic concepts, which is further developed in his book
“Origins of Life” published in 19856. He invoked Lynn Margulis’ ideas on the origin
of eukaryotic cells, proposing that the pre-biotic evolution would have been
accomplished by the independent formation, on one hand, of metabolic systems,
and on the other hand, of autoreplicative molecules. At a specific moment, some
of these molecules would have been synthesized within some of those systems,
first as parasites and then as symbionts, which evolved together. Under this
concept, the primitive nucleic acids, namely RNAs, invaded their metabolic hosts
and used them for their own replication, in a schema summarized “metabolism
first, replication second”7. Although, in these circumstances, the replication of
nucleic acids could be unlikely, it gives an interesting solution to the main
dilemma, how primitive systems could be able to storage information and to
perpetuate themselves, using nucleic acids. However, in terms of biochemical
principles, the replication process seems to be simpler than the metabolism one,
which leads us to the belief that this process could have had the leading role in
this evolutive scenario. But as Marcello Barbieri points out, “it remains to be seen,
however, if the replication paradigm can really account for the processes that led
to the origin of the first cells.”5.
Following the replication paradigm, the formulation of the so called “RNA world”
by Walter Gilbert in 1986 to designate a hypothetical stage in the development of
life in which RNA was the primary information and catalytic molecule, combining
the properties of RNA and proteins, was a natural theoretical step after the
discovery of ribozymes by Thomas Cech and Sidney Altman. The fact that
ribozymes came before protein enzymes does not mean that replication came
before metabolism, although it was generally interpreted that if RNAs could
behave as genes and enzymes, they became the first replicators in the history of
life5. According to this scenario the RNA world should be a world of replicators,
implying that primitive Earth was populated by RNAs showing the capacity to
replicate themselves or to catalyse the replication of other RNAs5. This has not
been proven yet. Furthermore, RNAs are “sophisticated, evolutionarily advanced
molecules”7,which could imply that the latter were not the main leading
molecules responsible for the chemical evolution on primitive Earth. In this sense,
we would like to think that a symbiogenic scenario was responsible for the
development of pre-biotic processes that led to the origin of primitive cells.
In his article “The Universal Ancestor”9, Carl Woese presents a genetic model for
the universal ancestor of extant life, pointing out that “the universal ancestor is
not a discrete entity. It is, rather, a diverse community of cells that survives and
evolves as a biological unit”. He named this universal ancestor progenote. We
believe that this theoretical scenario fits correctly into the idea of a cell
community that existed and evolved in the early life stages on Earth. On this
sense, the traditional concept of “primordial cell” is a biological chimera in our
point of view. We suggest there were probably open communities of protocells,
changing and incorporating information among them, as the base supporting
evolution of the early “tree” of life. Ultimately, we consider that primitive reality
to a huge “prebiotic web library”. From this point of view, one important question
must be formulated. Did those protocells evolve from elements present only on
Earth, or did they incorporate alien materials? Even if we still do not have a final
answer, both scenarios should always be presented in any symbiogenic strategy.
Cells, like we have previously described, can be included in a new cellular concept
entitled, “symbiocell”, since survival of the community under such adverse
conditions required a cooperative, synergistic strategy. A “symbiocell” is a
concept based on the principles that the cell or the protocell is organized not only
as a separate biological unit, but as a community of functions acquired by
symbiogenic means and evolved using symbiotic rules. In a sense, evolving and
behavioring like an ecological community or a natural microcosm. The
“symbiocell” will be the material support for the existence and development of
the adequate information flow, energy and metabolites that constitute the web
base of the autonomous and further communal life evolution. Later, these cells
continue to evolve in a symbiogenic way, involving and acquiring specialized
microbial communities which lead to the appearance of the eukaryotic cells,
according to the serial endosymbiotic theory10.
4. FINAL REMARKS
In this paper we have used a new approach to understand the problem of the
origin of life on Earth. According to this point of view, we believe that the
metabolism/replication dilemma is not the main issue to be considered. The main
point will be how evolution took place and worked in such early adverse
environment. To obtain a good answer to this problem, we must consider
different evolutive approaches, other than the traditional Darwinian ones,
introducing new tools to build a symbiogenic scenario of evolution. A cooperative
and a communal synergistic strategy should be considered as having been the
determinant in the development of the “survival of the fittest”, especially under
such extremely adverse environmental conditions. This approach could also be
used to understand the life formation in other locations beyond our planet.
Human evolution
Human evolution is the evolutionary process that led to the emergence of
anatomically modern humans, beginning with the evolutionary history of
primates—in particular genus Homo—and leading to the emergence of Homo
sapiens as a distinct species of the hominid family, the great apes. This process
involved the gradual development of traits such as human bipedalism and
language as well as interbreeding with other hominins, which indicate that
human evolution was not linear but a web.
Within the Hominoidea (apes) superfamily, the Hominidae family diverged from
the Hylobatidae (gibbon) family some 15–20 million years ago; African great apes
(subfamily Homininae) diverged from orangutans (Ponginae) about 14 million
years ago; the Hominini tribe (humans, Australopithecines and other extinct biped
genera, and chimpanzees) parted from the Gorillini tribe (gorillas) between 8–9
million years ago; and, in turn, the subtribes Hominina (humans and biped
ancestors) and Panina (chimpanzees) separated 4–7 million years ago
According to the recent African origin of modern humans theory, modern humans
evolved in Africa possibly from Homo heidelbergensis, Homo rhodesiensis or
Homo antecessor and migrated out of the continent some 50,000 to 100,000
years ago, gradually replacing local populations of Homo erectus, Denisova
hominins, Homo floresiensis, Homo luzonensis and Homo
neanderthalensis.Archaic Homo sapiens, the forerunner of anatomically modern
humans, evolved in the Middle Paleolithic between 400,000 and 250,000 years
ago.Recent DNA evidence suggests that several haplotypes of Neanderthal origin
are present among all non-African populations, and Neanderthals and other
hominins, such as Denisovans, may have contributed up to 6% of their genome to
present-day humans, suggestive of a limited interbreeding between these
species.The transition to behavioral modernity with the development of symbolic
culture, language, and specialized lithic technology happened around 50,000
years ago, according to some anthropologists] although others point to evidence
that suggests that a gradual change in behavior took place over a longer time
span.
Homo sapiens is the only extant species of its genus, Homo. While some (extinct)
Homo species might have been ancestors of Homo sapiens, many, perhaps most,
were likely "cousins", having speciated away from the ancestral hominin line.
There is yet no consensus as to which of these groups should be considered a
separate species and which should be a subspecies; this may be due to the dearth
of fossils or to the slight differences used to classify species in the genus
Homo.The Sahara pump theory (describing an occasionally passable "wet" Sahara
desert) provides one possible explanation of the early variation in the genus
Homo.
Based on archaeological and paleontological evidence, it has been possible to
infer, to some extent, the ancient dietary practices of various Homo species and
to study the role of diet in physical and behavioral evolution within Homo.
In Africa in the Early Pleistocene, 1.5–1 Ma, some populations of Homo habilis are
thought to have evolved larger brains and to have made more elaborate stone
tools; these differences and others are sufficient for anthropologists to classify
them as a new species, Homo erectus—in Africa.The evolution of locking knees
and the movement of the foramen magnum are thought to be likely drivers of the
larger population changes. This species also may have used fire to cook meat.
Richard Wrangham suggests that the fact that Homo seems to have been ground
dwelling, with reduced intestinal length, smaller dentition, "and swelled our
brains to their current, horrendously fuel-inefficient size" suggest that control of
fire and releasing increased nutritional value through cooking was the key
adaptation that separated Homo from tree-sleeping Australopithecines.