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Biology

For other uses, see Biology (disambiguation).


"Biological" redirects here. For other uses, see Biological
(disambiguation).

Biology is the scientific study of life.


[1][2][3] It is a natural science with a

broad scope but has several unifying


themes that tie it together as a single,
coherent field.[1][2][3] For instance, all
organisms are made up of cells that
process hereditary information
encoded in genes, which can be
transmitted to future generations.
Another major theme is evolution,
which explains the unity and diversity
of life.[1][2][3] Energy processing is
also important to life as it allows
organisms to move, grow, and
reproduce.[1][2][3] Finally, all
organisms are able to regulate their
own internal environments.[1][2][3][4][5]

Biologists are able to study life at


multiple levels of organization,[1] from Biology deals with the study of life.
the molecular biology of a cell to the top: E. coli bacteria and gazelle
anatomy and physiology of plants and bottom: Goliath beetle and tree
fern
animals, and evolution
of populations.[1][6] Part of a series on
Hence, there are Biology
multiple subdisciplines
:
within biology, each
defined by the nature of
their research questions
and the tools that they
use.[7][8][9] Like other Science of life
scientists, biologists
use the scientific Index
method to make Outline
observations, pose Glossary
questions, generate History (timeline)
hypotheses, perform
experiments, and form Key components
conclusions about the Cell theory
world around them. [1]
Ecosystem
Life on Earth, which Evolution
emerged more than 3.7 Phylogeny
billion years ago,[10] is
Properties of life
immensely diverse.
Biologists have sought Adaptation
to study and classify Energy processing
the various forms of life,
Growth
from prokaryotic
Order
organisms such as
archaea and bacteria to Regulation
eukaryotic organisms Reproduction
such as protists, fungi, Response to environment
plants, and animals.
Domains and Kingdoms of life
These various
organisms contribute to Archaea
the biodiversity of an Bacteria
ecosystem, where they
Eukarya (Animals, Fungi, Plants, Protists)
:
play specialized roles in
Branches
the cycling of nutrients
and energy through Anatomy

their biophysical Biotechnology


environment. Botany
Cell biology
Contents Ecology
1 Etymology Evolutionary biology
2 History Genetics
3 Fundamentals Marine biology
3.1 Chemical basis
Microbiology
3.1.1 Atoms and
Molecular biology
molecules
3.1.2 Water Mycology

3.1.3 Organic Neuroscience


compounds Paleontology
3.1.4 Macromolecules Phycology
3.2 Cells
Physiology
3.2.1 Cell structure
Protistology
3.2.2 Metabolism
3.2.3 Cellular Virology

respiration Zoology
3.2.4 Photosynthesis
3.2.5 Cell signaling Research
3.2.6 Cell cycle Biologist (list)
3.3 Genetics List of biology awards
3.3.1 Inheritance List of journals
3.3.2 DNA
List of research methods
3.3.3 Gene expression
List of unsolved problems
3.3.4 Gene regulation
3.3.5 Genomes
Applications
:
3.3.6 Biotechnology Agricultural science
3.3.7 Genes, Biomedical sciences
development, and Health technology
evolution
Pharming
3.4 Evolution
3.4.1 Evolutionary
processes Biology portal
3.4.2 Speciation
3.4.3 Phylogeny
3.4.4 History of life Category
3.5 Diversity
3.5.1 Bacteria and v
Archaea t
3.5.2 Protists e
3.5.3 Plant diversity
3.5.4 Fungi
3.5.5 Animal diversity
3.5.6 Viruses
3.6 Plant form and function
3.6.1 Plant body
3.6.2 Plant nutrition and transport
3.6.3 Plant development
3.6.4 Plant reproduction
3.6.5 Plant responses
3.7 Animal form and function
3.7.1 General features
3.7.2 Water and salt balance
3.7.3 Nutrition and digestion
3.7.4 Breathing
3.7.5 Circulation
3.7.6 Muscle and movement
3.7.7 Nervous system
:
3.7.8 Hormonal control
3.7.9 Animal reproduction
3.7.10 Animal development
3.7.11 Immune system
3.7.12 Animal behavior
3.8 Ecology
3.8.1 Ecosystems
3.8.2 Populations
3.8.3 Communities
3.8.4 Biosphere
3.8.5 Conservation
4 See also
5 References
6 Further reading
7 External links

Etymology
Biology derives from the Ancient Greek words of βίος romanized
bíos meaning 'life' and -λογία; romanized -logía meaning 'branch of
study' or 'to speak'.[11][12] Those combined make the Greek word
βιολογία romanized biología meaning 'biology'. Despite this, the term
βιολογία as a whole did not exist in Ancient Greek. The first to
borrow it was the English and French (biologie). Historically there
was another term for biology in English, lifelore; it is rarely used
today.

The Latin-language form of the term first appeared in 1736 when


Swedish scientist Carl Linnaeus (Carl von Linné) used biologi in his
Bibliotheca Botanica. It was used again in 1766 in a work entitled
Philosophiae naturalis sive physicae: tomus III, continens geologian,
biologian, phytologian generalis, by Michael Christoph Hanov, a
disciple of Christian Wolff. The first German use, Biologie, was in a
:
1771 translation of Linnaeus' work. In 1797, Theodor Georg August
Roose used the term in the preface of a book, Grundzüge der Lehre
van der Lebenskraft. Karl Friedrich Burdach used the term in 1800 in
a more restricted sense of the study of human beings from a
morphological, physiological and psychological perspective
(Propädeutik zum Studien der gesammten Heilkunst). The term
came into its modern usage with the six-volume treatise Biologie,
oder Philosophie der lebenden Natur (1802–22) by Gottfried
Reinhold Treviranus, who announced:[13]

The objects of our research will be the different forms and


manifestations of life, the conditions and laws under which
these phenomena occur, and the causes through which they
have been affected. The science that concerns itself with these
objects we will indicate by the name biology [Biologie] or the
doctrine of life [Lebenslehre].

Many other terms used in biology to describe plants, animals,


diseases, and drugs have been derived from Greek and Latin due to
the historical contributions of the Ancient Greek and Roman
civilizations as well as the continued use of these two languages in
European universities during the Middle Ages and at the beginning of
the Renaissance.[14]

History
Further information: History of biology

The earliest of roots of science, which included medicine, can be


traced to ancient Egypt and Mesopotamia in around 3000 to 1200
BCE.[15][16] Their contributions later entered and shaped Greek
natural philosophy of classical antiquity.[15][16][17][18] Ancient Greek
philosophers such as Aristotle (384–322 BCE) contributed
:
extensively to the development of
biological knowledge. His works such as
History of Animals were especially
important because they revealed his
naturalist leanings, and later more
empirical works that focused on biological
causation and the diversity of life.
Aristotle's successor at the Lyceum,
Theophrastus, wrote a series of books on
botany that survived as the most
important contribution of antiquity to the
plant sciences, even into the Middle Ages.
[19]
Diagram of a fly from Robert
Hooke's innovative Micrographia,
Scholars of the medieval Islamic world 1665
who wrote on biology included al-Jahiz
(781–869), Al-Dīnawarī (828–896), who wrote on botany,[20] and
Rhazes (865–925) who wrote on anatomy and physiology. Medicine
was especially well studied by Islamic scholars working in Greek
philosopher traditions, while natural history drew heavily on
Aristotelian thought, especially in upholding a fixed hierarchy of life.

Biology began to quickly develop and grow with Anton van


Leeuwenhoek's dramatic improvement of the microscope. It was
then that scholars discovered spermatozoa, bacteria, infusoria and
the diversity of microscopic life. Investigations by Jan Swammerdam
led to new interest in entomology and helped to develop the basic
techniques of microscopic dissection and staining.[21]

Advances in microscopy also had a profound impact on biological


thinking. In the early 19th century, a number of biologists pointed to
the central importance of the cell. Then, in 1838, Schleiden and
Schwann began promoting the now universal ideas that (1) the basic
:
unit of organisms is the cell and (2) that individual cells have all the
characteristics of life, although they opposed the idea that (3) all
cells come from the division of other cells. However, Robert Remak
and Rudolf Virchow were able to reify the third tenet, and by the
1860s most biologists accepted all three tenets which consolidated
into cell theory.[22][23]

Meanwhile, taxonomy and classification became the focus of natural


historians. Carl Linnaeus published a basic taxonomy for the natural
world in 1735 (variations of which have been in use ever since), and
in the 1750s introduced scientific names for all his species.[24]
Georges-Louis Leclerc, Comte de Buffon, treated species as artificial
categories and living forms as malleable—even suggesting the
possibility of common descent. Although he was opposed to
evolution, Buffon is a key figure in the history of evolutionary
thought; his work influenced the evolutionary theories of both
Lamarck and Darwin.[25]

Serious evolutionary thinking originated


with the works of Jean-Baptiste Lamarck,
who was the first to present a coherent
theory of evolution.[27] He posited that
evolution was the result of environmental
stress on properties of animals, meaning
that the more frequently and rigorously an
organ was used, the more complex and
efficient it would become, thus adapting
the animal to its environment. Lamarck
believed that these acquired traits could In 1842, Charles Darwin penned
his first sketch of On the Origin
then be passed on to the animal's of Species.[26]
offspring, who would further develop and
perfect them.[28] However, it was the British naturalist Charles
Darwin, combining the biogeographical approach of Humboldt, the
:
uniformitarian geology of Lyell, Malthus's writings on population
growth, and his own morphological expertise and extensive natural
observations, who forged a more successful evolutionary theory
based on natural selection; similar reasoning and evidence led Alfred
Russel Wallace to independently reach the same conclusions.[29][30]
Darwin's theory of evolution by natural selection quickly spread
through the scientific community and soon became a central axiom
of the rapidly developing science of biology.

The basis for modern genetics began with the work of Gregor
Mendel, who presented his paper, "Versuche über
Pflanzenhybriden" ("Experiments on Plant Hybridization"), in 1865,
[31] which outlined the principles of biological inheritance, serving as

the basis for modern genetics.[32] However, the significance of his


work was not realized until the early 20th century when evolution
became a unified theory as the modern synthesis reconciled
Darwinian evolution with classical genetics.[33] In the 1940s and
early 1950s, a series of experiments by Alfred Hershey and Martha
Chase pointed to DNA as the component of chromosomes that held
the trait-carrying units that had become known as genes. A focus on
new kinds of model organisms such as viruses and bacteria, along
with the discovery of the double-helical structure of DNA by James
Watson and Francis Crick in 1953, marked the transition to the era of
molecular genetics. From the 1950s onwards, biology has been
vastly extended in the molecular domain. The genetic code was
cracked by Har Gobind Khorana, Robert W. Holley and Marshall
Warren Nirenberg after DNA was understood to contain codons.
Finally, the Human Genome Project was launched in 1990 with the
goal of mapping the general human genome. This project was
essentially completed in 2003,[34] with further analysis still being
published. The Human Genome Project was the first step in a
globalized effort to incorporate accumulated knowledge of biology
:
into a functional, molecular definition of the human body and the
bodies of other organisms.

Fundamentals
Chemical basis

Atoms and molecules

Further information: Chemistry

All organisms are made up of matter and


all matter is made up of elements.[35]
Oxygen, carbon, hydrogen, and nitrogen
are the four elements that account for
96% of all organisms, with calcium,
phosphorus, sulfur, sodium, chlorine, and In the Bohr model of an atom,
magnesium constituting the remaining electrons (blue dot) orbit around
3.7%.[35] Different elements can combine an atomic nucleus (red-filled
circle) in specific atomic orbitals
to form compounds such as water, which (grey empty circles).
is fundamental to life.[35] Life on Earth
began from water and remained there for about three billions years
prior to migrating onto land.[36] Matter can exist in different states as
a solid, liquid, or gas.

The smallest unit of an element is an atom, which is composed of an


atomic nucleus and one or more electrons moving around the
nucleus, as described by the Bohr model.[37] The nucleus is made of
one or more protons and a number of neutrons. Protons have a
positive electric charge, neutrons are electrically neutral, and
electrons have a negative electric charge.[38] Atoms with equal
numbers of protons and electrons are electrically neutral. The atom
of each specific element contains a unique number of protons, which
:
is known as its atomic number, and the sum of its protons and
neutrons is an atom's mass number. The masses of individual
protons, neutrons, and electrons can be measured in grams or
Daltons (Da), with the mass of each proton or neutron rounded to 1
Da.[38] Although all atoms of a specific element have the same
number of protons, they may differ in the number of neutrons,
thereby existing as isotopes.[35] Carbon, for example, can exist as a
stable isotope (carbon-12 or carbon-13) or as a radioactive isotope
(carbon-14), the latter of which can be used in radiometric dating
(specifically radiocarbon dating) to determine the age of organic
materials.[35]

Individual atoms can be held together by chemical bonds to form


molecules and ionic compounds.[35] Common types of chemical
bonds include ionic bonds, covalent bonds, and hydrogen bonds.
Ionic bonding involves the electrostatic attraction between
oppositely charged ions, or between two atoms with sharply
different electronegativities,[39] and is the primary interaction
occurring in ionic compounds. Ions are atoms (or groups of atoms)
with an electrostatic charge. Atoms that gain electrons make
negatively charged ions (called anions) whereas those that lose
electrons make positively charged ions (called cations).

Unlike ionic bonds, a covalent bond involves the sharing of electron


pairs between atoms. These electron pairs and the stable balance of
attractive and repulsive forces between atoms, when they share
electrons, is known as covalent bonding.[40][better source needed]

A hydrogen bond is primarily an electrostatic force of attraction


between a hydrogen atom which is covalently bound to a more
electronegative atom or group such as oxygen. A ubiquitous example
of a hydrogen bond is found between water molecules. In a discrete
water molecule, there are two hydrogen atoms and one oxygen atom.
:
Two molecules of water can form a hydrogen bond between them.
When more molecules are present, as is the case with liquid water,
more bonds are possible because the oxygen of one water molecule
has two lone pairs of electrons, each of which can form a hydrogen
bond with a hydrogen on another water molecule.

Water

Life arose from the Earth's first ocean,


which was formed approximately 3.8
billion years ago.[38] Since then, water
continues to be the most abundant
molecule in every organism. Water is
important to life because it is an effective
solvent, capable of dissolving solutes such
as sodium and chloride ions or other small
molecules to form an aqueous solution. Model of hydrogen bonds (1)
Once dissolved in water, these solutes are between molecules of water
more likely to come in contact with one another and therefore take
part in chemical reactions that sustain life.[38]

In terms of its molecular structure, water is a small polar molecule


with a bent shape formed by the polar covalent bonds of two
hydrogen (H) atoms to one oxygen (O) atom (H2O).[38] Because the
O–H bonds are polar, the oxygen atom has a slight negative charge
and the two hydrogen atoms have a slight positive charge.[38] This
polar property of water allows it to attract other water molecules via
hydrogen bonds, which makes water cohesive.[38] Surface tension
results from the cohesive force due to the attraction between
molecules at the surface of the liquid.[38] Water is also adhesive as it
is able to adhere to the surface of any polar or charged non-water
molecules.[38]
:
Water is denser as a liquid than it is as a solid (or ice).[38] This unique
property of water allows ice to float above liquid water such as
ponds, lakes, and oceans, thereby insulating the liquid below from
the cold air above.[38] The lower density of ice compared to liquid
water is due to the lower number of water molecules that form the
crystal lattice structure of ice, which leaves a large amount of space
between water molecules.[38] In contrast, there is no crystal lattice
structure in liquid water, which allows more water molecules to
occupy the same amount of volume.[38]

Water also has the capacity to absorb energy, giving it a higher


specific heat capacity than other solvents such as ethanol.[38] Thus,
a large amount of energy is needed to break the hydrogen bonds
between water molecules to convert liquid water into gas (or water
vapor).[38]

As a molecule, water is not completely stable as each water molecule


continuously dissociates into hydrogen and hydroxyl ions before
reforming into a water molecule again.[38] In pure water, the number
of hydrogen ions balances (or equals) the number of hydroxyl ions,
resulting in a pH that is neutral. If hydrogen ions were to exceed
hydroxyl ions, then the pH of the solution would be acidic.
Conversely, a solution's pH would turn basic if hydroxyl ions were to
exceed hydrogen ions.

Organic compounds

Further information: Organic chemistry

Organic compounds are molecules that contain carbon bonded to


another element such as hydrogen.[38] With the exception of water,
nearly all the molecules that make up each organism contain carbon.
[38][41] Carbon has six electrons, two of which are located in its first

shell, leaving four electrons in its valence shell. Thus, carbon can
:
form covalent bonds with up to four other
atoms, making it the most versatile atom
on Earth as it is able to form diverse, large,
and complex molecules.[38][41] For
example, a single carbon atom can form
four single covalent bonds such as in
methane, two double covalent bonds such
Organic compounds such as
as in carbon dioxide (CO2), or a triple glucose are vital to organisms.
covalent bond such as in carbon monoxide
(CO). Moreover, carbon can form very long chains of interconnecting
carbon–carbon bonds such as octane or ring-like structures such as
glucose.

The simplest form of an organic molecule is the hydrocarbon, which


is a large family of organic compounds that are composed of
hydrogen atoms bonded to a chain of carbon atoms. A hydrocarbon
backbone can be substituted by other elements such as oxygen (O),
hydrogen (H), phosphorus (P), and sulfur (S), which can change the
chemical behavior of that compound.[38] Groups of atoms that
contain these elements (O-, H-, P-, and S-) and are bonded to a
central carbon atom or skeleton are called functional groups.[38]
There are six prominent functional groups that can be found in
organisms: amino group, carboxyl group, carbonyl group, hydroxyl
group, phosphate group, and sulfhydryl group.[38]

In 1953, Stanley Miller and Harold Urey conducted a classic


experiment (otherwise known as the Miller-Urey experiment), which
showed that organic compounds could be synthesized abiotically
within a closed system that mimicked the conditions of early Earth,
leading them to conclude that complex organic molecules could
have arisen spontaneously in early Earth, most likely near volcanoes,
and could have part of the early stages of abiogenesis (or origin of
life).[42][38]
:
Macromolecules

Further information: Biochemistry

Macromolecules are large molecules made


up of smaller molecular subunits that are
joined together.[43] Small molecules such
as sugars, amino acids, and nucleotides
can act as single repeating units called
monomers to form chain-like molecules A phospholipid bilayer consists of
two adjacent sheets of
called polymers via a chemical process
phospholipids, with the
called condensation. [44] For example, hydrophilic tails facing inwards
amino acids can form polypeptides and the hydrophobic heads
facing outwards.
whereas nucleotides can form strands of
nucleic acid. Polymers make up three of the four macromolecules
(polysaccharides, lipids, proteins, and nucleic acids) that are found in
all organisms. Each of these macromolecules plays a specialized role
within any given cell.

Carbohydrates (or sugar) are molecules with the molecular formula


(CH2O)n, with n being the number of carbon-hydrate groups.[45]
They include monosaccharides (monomer), oligosaccharides (small
polymers), and polysaccharides (large polymers). Monosaccharides
can be linked together by glycosidic linkages, a type of covalent
bond.[45] When two monosaccharides such as glucose and fructose
are linked together, they can form a disaccharide such as sucrose.
[45] When many monosaccharides are linked together, they can form

an oligosaccharide or a polysaccharide, depending on the number of


monosaccharides. Polysaccharides can vary in function.
Monosaccharides such as glucose can be a source of energy and
some polysaccharides can serve as storage material that can be
hydrolyzed to provide cells with sugar.
:
Lipids are the only class of macromolecules that are not made up of
polymers. The most biologically important lipids are steroids,
phospholipids, and fats.[44] These lipids are organic compounds that
are largely nonpolar and hydrophobic.[46] Steroids are organic
compounds that consist of four fused rings.[46] Phospholipids
consist of glycerol that is linked to a phosphate group and two
hydrocarbon chains (or fatty acids).[46] The glycerol and phosphate
group together constitute the polar and hydrophilic (or head) region
of the molecule whereas the fatty acids make up the nonpolar and
hydrophobic (or tail) region.[46] Thus, when in water, phospholipids
tend to form a phospholipid bilayer whereby the hydrophobic heads
face outwards to interact with water molecules. Conversely, the
hydrophobic tails face inwards towards other hydrophobic tails to
avoid contact with water.[46]

Proteins are the most diverse of the


macromolecules, which include enzymes,
transport proteins, large signaling
molecules, antibodies, and structural
proteins. The basic unit (or monomer) of a
protein is an amino acid, which has a
central carbon atom that is covalently
bonded to a hydrogen atom, an amino The (a) primary, (b) secondary,
group, a carboxyl group, and a side chain (c) tertiary, and (d) quaternary
structures of a hemoglobin
(or R-group, "R" for residue).[43] There are protein
twenty amino acids that make up the
building blocks of proteins, with each amino acid having its own
unique side chain.[43] The polarity and charge of the side chains
affect the solubility of amino acids. An amino acid with a side chain
that is polar and electrically charged is soluble as it is hydrophilic
whereas an amino acid with a side chain that lacks a charged or an
electronegative atom is hydrophobic and therefore tends to coalesce
:
rather than dissolve in water.[43] Proteins have four distinct levels of
organization (primary, secondary, tertiary, and quartenary). The
primary structure consists of a unique sequence of amino acids that
are covalently linked together by peptide bonds.[43] The side chains
of the individual amino acids can then interact with each other, giving
rise to the secondary structure of a protein.[43] The two common
types of secondary structures are alpha helices and beta sheets.[43]
The folding of alpha helices and beta sheets gives a protein its three-
dimensional or tertiary structure. Finally, multiple tertiary structures
can combine to form the quaternary structure of a protein.

Nucleic acids are polymers made up of monomers called


nucleotides.[47] Their function is to store, transmit, and express
hereditary information.[44] Nucleotides consist of a phosphate group,
a five-carbon sugar, and a nitrogenous base. Ribonucleotides, which
contain ribose as the sugar, are the monomers of ribonucleic acid
(RNA). In contrast, deoxyribonucleotides contain deoxyribose as the
sugar and are constitute the monomers of deoxyribonucleic acid
(DNA). RNA and DNA also differ with respect to one of their bases.
[47] There are two types of bases: purines and pyrimidines.[47] The

purines include guanine (G) and adenine (A) whereas the pyrimidines
consist of cytosine (C), uracil (U), and thymine (T). Uracil is used in
RNA whereas thymine is used in DNA. Taken together, when the
different sugar and bases are take into consideration, there are eight
distinct nucleotides that can form two types of nucleic acids: DNA
(A, G, C, and T) and RNA (A, G, C, and U).[47]

Cells

Further information: Cell biology

Cell theory states that cells are the fundamental units of life, that all
living things are composed of one or more cells, and that all cells
:
arise from preexisting cells through cell division.[48] Most cells are
very small, with diameters ranging from 1 to 100 micrometers and are
therefore only visible under a light or electron microscope.[49] There
are generally two types of cells: eukaryotic cells, which contain a
nucleus, and prokaryotic cells, which do not. Prokaryotes are single-
celled organisms such as bacteria, whereas eukaryotes can be
single-celled or multicellular. In multicellular organisms, every cell in
the organism's body is derived ultimately from a single cell in a
fertilized egg.

Cell structure

Every cell is enclosed within a cell


membrane that separates its cytoplasm
from the extracellular space.[50] A cell
membrane consists of a lipid bilayer,
including cholesterols that sit between
phospholipids to maintain their fluidity at Structure of an animal cell
various temperatures. Cell membranes are depicting various organelles
semipermeable, allowing small molecules such as oxygen, carbon
dioxide, and water to pass through while restricting the movement of
larger molecules and charged particles such as ions.[51] Cell
membranes also contains membrane proteins, including integral
membrane proteins that go across the membrane serving as
membrane transporters, and peripheral proteins that loosely attach
to the outer side of the cell membrane, acting as enzymes shaping
the cell.[52] Cell membranes are involved in various cellular
processes such as cell adhesion, storing electrical energy, and cell
signalling and serve as the attachment surface for several
extracellular structures such as a cell wall, glycocalyx, and
cytoskeleton.

Within the cytoplasm of a cell, there are many biomolecules such as


:
proteins and nucleic acids.[53] In addition
to biomolecules, eukaryotic cells have
specialized structures called organelles
that have their own lipid bilayers or are
spatially units.[54] These organelles
include the cell nucleus, which contains
Structure of a plant cell most of the cell's DNA, or mitochondria,
which generates adenosine triphosphate
(ATP) to power cellular processes. Other organelles such as
endoplasmic reticulum and Golgi apparatus play a role in the
synthesis and packaging of proteins, respectively. Biomolecules
such as proteins can be engulfed by lysosomes, another specialized
organelle. Plant cells have additional organelles that distinguish them
from animal cells such as a cell wall that provides support for the
plant cell, chloroplasts that harvest sunlight energy to produce sugar,
and vacuoles that provide storage and structural support as well as
being involved in reproduction and breakdown of plant seeds.[54]
Eukaryotic cells also have cytoskeleton that is made up of
microtubules, intermediate filaments, and microfilaments, all of
which provide support for the cell and are involved in the movement
of the cell and its organelles.[54] In terms of their structural
composition, the microtubules are made up of tubulin (e.g., α-tubulin
and β-tubulin whereas intermediate filaments are made up of fibrous
proteins.[54] Microfilaments are made up of actin molecules that
interact with other strands of proteins.[54]

Metabolism

Further information: Bioenergetics

All cells require energy to sustain cellular processes. Energy is the


capacity to do work, which, in thermodynamics, can be calculated
using Gibbs free energy. According to the first law of
:
thermodynamics, energy is conserved, i.e.,
cannot be created or destroyed. Hence,
chemical reactions in a cell do not create
new energy but are involved instead in the
transformation and transfer of energy.[55]
Nevertheless, all energy transfers lead to
some loss of usable energy, which
Example of an enzyme-catalysed
increases entropy (or state of disorder) as exothermic reaction
stated by the second law of
thermodynamics. As a result, an organism requires continuous input
of energy to maintain a low state of entropy. In cells, energy can be
transferred as electrons during redox (reduction–oxidation)
reactions, stored in covalent bonds, and generated by the movement
of ions (e.g., hydrogen, sodium, potassium) across a membrane.

Metabolism is the set of life-sustaining chemical reactions in


organisms. The three main purposes of metabolism are: the
conversion of food to energy to run cellular processes; the
conversion of food/fuel to building blocks for proteins, lipids, nucleic
acids, and some carbohydrates; and the elimination of metabolic
wastes. These enzyme-catalyzed reactions allow organisms to grow
and reproduce, maintain their structures, and respond to their
environments. Metabolic reactions may be categorized as catabolic
—the breaking down of compounds (for example, the breaking down
of glucose to pyruvate by cellular respiration); or anabolic—the
building up (synthesis) of compounds (such as proteins,
carbohydrates, lipids, and nucleic acids). Usually, catabolism
releases energy, and anabolism consumes energy.

The chemical reactions of metabolism are organized into metabolic


pathways, in which one chemical is transformed through a series of
steps into another chemical, each step being facilitated by a specific
enzyme. Enzymes are crucial to metabolism because they allow
:
organisms to drive desirable reactions that require energy that will
not occur by themselves, by coupling them to spontaneous reactions
that release energy. Enzymes act as catalysts—they allow a reaction
to proceed more rapidly without being consumed by it—by reducing
the amount of activation energy needed to convert reactants into
products. Enzymes also allow the regulation of the rate of a
metabolic reaction, for example in response to changes in the cell's
environment or to signals from other cells.

Cellular respiration

Cellular respiration is a set of


metabolic reactions and
processes that take place in the
cells of organisms to convert
chemical energy from nutrients
into adenosine triphosphate
(ATP), and then release waste
products.[56] The reactions
involved in respiration are Respiration in a eukaryotic cell

catabolic reactions, which break


large molecules into smaller ones, releasing energy because weak
high-energy bonds, in particular in molecular oxygen,[57] are
replaced by stronger bonds in the products. Respiration is one of the
key ways a cell releases chemical energy to fuel cellular activity. The
overall reaction occurs in a series of biochemical steps, some of
which are redox reactions. Although cellular respiration is technically
a combustion reaction, it clearly does not resemble one when it
occurs in a cell because of the slow, controlled release of energy
from the series of reactions.

Sugar in the form of glucose is the main nutrient used by animal and
plant cells in respiration. Cellular respiration involving oxygen is
:
called aerobic respiration, which has four stages: glycolysis, citric
acid cycle (or Krebs cycle), electron transport chain, and oxidative
phosphorylation.[58] Glycolysis is a metabolic process that occurs in
the cytoplasm whereby glucose is converted into two pyruvates, with
two net molecules of ATP being produced at the same time.[58] Each
pyruvate is then oxidized into acetyl-CoA by the pyruvate
dehydrogenase complex, which also generates NADH and carbon
dioxide. Acetyl-Coa enters the citric acid cycle, which takes places
inside the mitochondrial matrix. At the end of the cycle, the total
yield from 1 glucose (or 2 pyruvates) is 6 NADH, 2 FADH2, and 2 ATP
molecules. Finally, the next stage is oxidative phosphorylation, which
in eukaryotes, occurs in the mitochondrial cristae. Oxidative
phosphorylation comprises the electron transport chain, which is a
series of four protein complexes that transfer electrons from one
complex to another, thereby releasing energy from NADH and FADH2
that is coupled to the pumping of protons (hydrogen ions) across the
inner mitochondrial membrane (chemiosmosis), which generates a
proton motive force.[58] Energy from the proton motive force drives
the enzyme ATP synthase to synthesize more ATPs by
phosphorylating ADPs. The transfer of electrons terminates with
molecular oxygen being the final electron acceptor.

If oxygen were not present, pyruvate would not be metabolized by


cellular respiration but undergoes a process of fermentation. The
pyruvate is not transported into the mitochondrion but remains in the
cytoplasm, where it is converted to waste products that may be
removed from the cell. This serves the purpose of oxidizing the
electron carriers so that they can perform glycolysis again and
removing the excess pyruvate. Fermentation oxidizes NADH to NAD+
so it can be re-used in glycolysis. In the absence of oxygen,
fermentation prevents the buildup of NADH in the cytoplasm and
provides NAD+ for glycolysis. This waste product varies depending
:
on the organism. In skeletal muscles, the waste product is lactic acid.
This type of fermentation is called lactic acid fermentation. In
strenuous exercise, when energy demands exceed energy supply,
the respiratory chain cannot process all of the hydrogen atoms
joined by NADH. During anaerobic glycolysis, NAD+ regenerates
when pairs of hydrogen combine with pyruvate to form lactate.
Lactate formation is catalyzed by lactate dehydrogenase in a
reversible reaction. Lactate can also be used as an indirect precursor
for liver glycogen. During recovery, when oxygen becomes available,
NAD+ attaches to hydrogen from lactate to form ATP. In yeast, the
waste products are ethanol and carbon dioxide. This type of
fermentation is known as alcoholic or ethanol fermentation. The ATP
generated in this process is made by substrate-level
phosphorylation, which does not require oxygen.

Photosynthesis

Photosynthesis is a process used by


plants and other organisms to convert light
energy into chemical energy that can later
be released to fuel the organism's
metabolic activities via cellular respiration.
This chemical energy is stored in
carbohydrate molecules, such as sugars,
which are synthesized from carbon dioxide
and water.[59][60][61] In most cases,
oxygen is also released as a waste
product. Most plants, algae, and Photosynthesis changes sunlight
into chemical energy, splits water
cyanobacteria perform photosynthesis,
to liberate O2, and fixes CO2 into
which is largely responsible for producing sugar.
and maintaining the oxygen content of the
Earth's atmosphere, and supplies most of the energy necessary for
life on Earth.[62]
:
Photosynthesis has four stages: Light absorption, electron transport,
ATP synthesis, and carbon fixation.[58] Light absorption is the initial
step of photosynthesis whereby light energy is absorbed by
chlorophyll pigments attached to proteins in the thylakoid
membranes. The absorbed light energy is used to remove electrons
from a donor (water) to a primary electron acceptor, a quinone
designated as Q. In the second stage, electrons move from the
quinone primary electron acceptor through a series of electron
carriers until they reach a final electron acceptor, which is usually the
oxidized form of NADP+, which is reduced to NADPH, a process that
takes place in a protein complex called photosystem I (PSI). The
transport of electrons is coupled to the movement of protons (or
hydrogen) from the stroma to the thylakoid membrane, which forms
a pH gradient across the membrane as hydrogen becomes more
concentrated in the lumen than in the stroma. This is analogous to
the proton-motive force generated across the inner mitochondrial
membrane in aerobic respiration.[58]

During the third stage of photosynthesis, the movement of protons


down their concentration gradients from the thylakoid lumen to the
stroma through the ATP synthase is coupled to the synthesis of ATP
by that same ATP synthase.[58] The NADPH and ATPs generated by
the light-dependent reactions in the second and third stages,
respectively, provide the energy and electrons to drive the synthesis
of glucose by fixing atmospheric carbon dioxide into existing organic
carbon compounds, such as ribulose bisphosphate (RuBP) in a
sequence of light-independent (or dark) reactions called the Calvin
cycle.[63]

Cell signaling

Cell communication (or signaling) is the ability of cells to receive,


process, and transmit signals with its environment and with itself.[64]
:
[65]Signals can be non-chemical such as light, electrical impulses,
and heat, or chemical signals (or ligands) that interact with
receptors, which can be found embedded in the cell membrane of
another cell or located deep inside a cell.[66][65] There are generally
four types of chemical signals: autocrine, paracrine, juxtacrine, and
hormones.[66] In autocrine signaling, the ligand affects the same cell
that releases it. Tumor cells, for example, can reproduce
uncontrollably because they release signals that initiate their own
self-division. In paracrine signaling, the ligand diffuses to nearby
cells and affect them. For example, brain cells called neurons release
ligands called neurotransmitters that diffuse across a synaptic cleft
to bind with a receptor on an adjacent cell such as another neuron or
muscle cell. In juxtacrine signaling, there is direct contact between
the signaling and responding cells. Finally, hormones are ligands that
travel through the circulatory systems of animals or vascular systems
of plants to reach their target cells. Once a ligand binds with a
receptor, it can influence the behavior of another cell, depending on
the type of receptor. For instance, neurotransmitters that bind with
an inotropic receptor can alter the excitability of a target cell. Other
types of receptors include protein kinase receptors (e.g., receptor for
the hormone insulin) and G protein-coupled receptors. Activation of
G protein-coupled receptors can initiate second messenger
cascades. The process by which a chemical or physical signal is
transmitted through a cell as a series of molecular events is called
signal transduction

Cell cycle

The cell cycle is a series of events that take place in a cell that cause
it to divide into two daughter cells. These events include the
duplication of its DNA and some of its organelles, and the
subsequent partitioning of its cytoplasm into two daughter cells in a
process called cell division.[67] In eukaryotes (i.e., animal, plant,
:
fungal, and protist cells), there
are two distinct types of cell
division: mitosis and meiosis.[68]
Mitosis is part of the cell cycle, in
which replicated chromosomes
are separated into two new
nuclei. Cell division gives rise to
genetically identical cells in In meiosis, the chromosomes duplicate and
the homologous chromosomes exchange
which the total number of
genetic information during meiosis I. The
chromosomes is maintained. In daughter cells divide again in meiosis II to
general, mitosis (division of the form haploid gametes.
nucleus) is preceded by the S
stage of interphase (during which the DNA is replicated) and is often
followed by telophase and cytokinesis; which divides the cytoplasm,
organelles and cell membrane of one cell into two new cells
containing roughly equal shares of these cellular components. The
different stages of mitosis all together define the mitotic phase of an
animal cell cycle—the division of the mother cell into two genetically
identical daughter cells.[69] The cell cycle is a vital process by which
a single-celled fertilized egg develops into a mature organism, as
well as the process by which hair, skin, blood cells, and some internal
organs are renewed. After cell division, each of the daughter cells
begin the interphase of a new cycle. In contrast to mitosis, meiosis
results in four haploid daughter cells by undergoing one round of
DNA replication followed by two divisions.[70] Homologous
chromosomes are separated in the first division (meiosis I), and
sister chromatids are separated in the second division (meiosis II).
Both of these cell division cycles are used in the process of sexual
reproduction at some point in their life cycle. Both are believed to be
present in the last eukaryotic common ancestor.

Prokaryotes (i.e., archaea and bacteria) can also undergo cell


:
division (or binary fission). Unlike the processes of mitosis and
meiosis in eukaryotes, binary fission takes in prokaryotes takes place
without the formation of a spindle apparatus on the cell. Before
binary fission, DNA in the bacterium is tightly coiled. After it has
uncoiled and duplicated, it is pulled to the separate poles of the
bacterium as it increases the size to prepare for splitting. Growth of a
new cell wall begins to separate the bacterium (triggered by FtsZ
polymerization and "Z-ring" formation)[71] The new cell wall
(septum) fully develops, resulting in the complete split of the
bacterium. The new daughter cells have tightly coiled DNA rods,
ribosomes, and plasmids.

Genetics

Inheritance

Further information: Classical genetics

Genetics is the scientific study of


inheritance.[72][73][74] Mendelian
inheritance, specifically, is the process by
which genes and traits are passed on from
parents to offspring.[32] It was formulated
by Gregor Mendel, based on his work with
pea plants in the mid-nineteenth century.
Mendel established several principles of
Punnett square depicting a cross
inheritance. The first is that genetic between two pea plants
characteristics, which are now called heterozygous for purple (B) and
white (b) blossoms
alleles, are discrete and have alternate
forms (e.g., purple vs. white or tall vs. dwarf), each inherited from
one of two parents. Based on his law of dominance and uniformity,
which states that some alleles are dominant while others are
recessive; an organism with at least one dominant allele will display
:
the phenotype of that dominant allele.[75] Exceptions to this rule
include penetrance and expressivity.[32] Mendel noted that during
gamete formation, the alleles for each gene segregate from each
other so that each gamete carries only one allele for each gene,
which is stated by his law of segregation. Heterozygotic individuals
produce gametes with an equal frequency of two alleles. Finally,
Mendel formulated the law of independent assortment, which states
that genes of different traits can segregate independently during the
formation of gametes, i.e., genes are unlinked. An exception to this
rule would include traits that are sex-linked. Test crosses can be
performed to experimentally determine the underlying genotype of
an organism with a dominant phenotype.[76] A Punnett square can
be used to predict the results of a test cross. The chromosome
theory of inheritance, which states that genes are found on
chromosomes, was supported by Thomas Morgans's experiments
with fruit flies, which established the sex linkage between eye color
and sex in these insects.[77] In humans and other mammals (e.g.,
dogs), it is not feasible or practical to conduct test cross
experiments. Instead, pedigrees, which are genetic representations
of family trees,[78] are used instead to trace the inheritance of a
specific trait or disease through multiple generations.[79]

DNA

A gene is a unit of heredity that corresponds to a region of


deoxyribonucleic acid (DNA) that carries genetic information that
influences the form or function of an organism in specific ways. DNA
is a molecule composed of two polynucleotide chains that coil
around each other to form a double helix, which was first described
by James Watson and Francis Crick in 1953.[80] It is found as linear
chromosomes in eukaryotes, and circular chromosomes in
prokaryotes. A chromosome is an organized structure consisting of
DNA and histones. The set of chromosomes in a cell and any other
:
hereditary information found in the
mitochondria, chloroplasts, or other locations is
collectively known as a cell's genome. In
eukaryotes, genomic DNA is localized in the
cell nucleus, or with small amounts in
mitochondria and chloroplasts.[81] In
prokaryotes, the DNA is held within an
irregularly shaped body in the cytoplasm called
the nucleoid.[82] The genetic information in a
genome is held within genes, and the complete
assemblage of this information in an organism
is called its genotype.[83] Genes encode the
Bases lie between two
information needed by cells for the synthesis ofspiraling DNA strands.
proteins, which in turn play a central role in
influencing the final phenotype of the organism.

The two polynucleotide strands that make up DNA run in opposite


directions to each other and are thus antiparallel. Each strand is
composed of nucleotides,[84][85] with each nucleotide containing
one of four nitrogenous bases (cytosine [C], guanine [G], adenine
[A] or thymine [T]), a sugar called deoxyribose, and a phosphate
group. The nucleotides are joined to one another in a chain by
covalent bonds between the sugar of one nucleotide and the
phosphate of the next, resulting in an alternating sugar-phosphate
backbone. It is the sequence of these four bases along the backbone
that encodes genetic information. Bases of the two polynucleotide
strands are bound together by hydrogen bonds, according to base
pairing rules (A with T and C with G), to make double-stranded DNA.
The bases are divided into two groups: pyrimidines and purines. In
DNA, the pyrimidines are thymine and cytosine whereas the purines
are adenine and guanine.

There are grooves that run along the entire length of the double helix
:
due to the uneven spacing of the DNA strands relative to each other.
[80] Both grooves differ in size, with the major groove being larger

and therefore more accessible to the binding of proteins than the


minor groove.[80] The outer edges of the bases are exposed to these
grooves and are therefore accessible for additional hydrogen
bonding.[80] Because each groove can have two possible base-pair
configurations (G-C and A-T), there are four possible base-pair
configurations within the entire double helix, each of which is
chemically distinct from another.[80] As a result, protein molecules
are able to recognize and bind to specific base-pair sequences,
which is the basis of specific DNA-protein interactions.

DNA replication is a semiconservative process whereby each strand


serves as a template for a new strand of DNA.[80] The process
begins with the unwounding of the double helix at an origin of
replication, which separates the two strands, thereby making them
available as two templates. This is then followed by the binding of
the enzyme primase to the template to synthesize a starter RNA (or
DNA in some viruses) strand called a primer from the 5’ to 3’
location.[80] Once the primer is completed, the primase is released
from the template, followed by the binding of the enzyme DNA
polymerase to the same template to synthesize new DNA. The rate
of DNA replication in a living cell was measured as 749 nucleotides
added per second under ideal conditions.[86]

DNA replication is not perfect as the DNA polymerase sometimes


insert bases that are not complementary to the template (e.g.,
putting in A in the strand opposite to G in the template strand).[80] In
eukaryotes, the initial error or mutation rate is about 1 in 100,000.[80]
Proofreading and mismatch repair are the two mechanisms that
repair these errors, which reduces the mutation rate to 10−10,
particularly before and after a cell cycle.[80]
:
Mutations are heritable changes in DNA.[80] They can arise
spontaneously as a result of replication errors that were not
corrected by proofreading or can be induced by an environmental
mutagen such as a chemical (e.g., nitrous acid, benzopyrene) or
radiation (e.g., x-ray, gamma ray, ultraviolet radiation, particles
emitted by unstable isotopes).[80] Mutations can appear as a change
in single base or at a larger scale involving chromosomal mutations
such as deletions, inversions, or translocations.[80]

In multicellular organisms, mutations can occur in somatic or


germline cells.[80] In somatic cells, the mutations are passed on to
daughter cells during mitosis.[80] In a germline cell such as a sperm
or an egg, the mutation will appear in an organism at fertilization.[80]
Mutations can lead to several types of phenotypic effects such as
silent, loss-of-function, gain-of-function, and conditional mutations.
[80]

Some mutations can be beneficial, as they are a source of genetic


variation for evolution.[80] Others can be harmful if they were to
result in a loss of function of genes needed for survival.[80]
Mutagens such as carcinogens are typically avoided as a matter of
public health policy goals.[80] One example is the banning of
chlorofluorocarbons (CFC) by the Montreal Protocol, as CFCs tend to
deplete the ozone layer, resulting in more ultraviolet radiation from
the sun passing through the Earth's upper atmosphere, thereby
causing somatic mutations that can lead to skin cancer.[80] Similarly,
smoking bans have been enforced throughout the world in an effort
to reduce the incidence of lung cancer.[80]

Gene expression

Gene expression is the molecular process by which a genotype gives


rise to a phenotype, i.e., observable trait. The genetic information
stored in DNA represents the genotype, whereas the phenotype
:
results from the synthesis of
proteins that control an
organism's structure and
development, or that act as
enzymes catalyzing specific
metabolic pathways. This
process is summarized by the The extended central dogma of molecular
biology includes all the processes involved in
central dogma of molecular the flow of genetic information.
biology, which was formulated by
Francis Crick in 1958.[87][88][89] According to the Central Dogma,
genetic information flows from DNA to RNA to protein. Hence, there
are two gene expression processes: transcription (DNA to RNA) and
translation (RNA to protein).[90] These processes are used by all life
—eukaryotes (including multicellular organisms), prokaryotes
(bacteria and archaea), and are exploited by viruses—to generate the
macromolecular machinery for life.

During transcription, messenger RNA (mRNA) strands are created


using DNA strands as a template, which is initiated when RNA
polymerase binds to a DNA sequence called a promoter, which
instructs the RNA to begin transcription of one of the two DNA
strands.[91] The DNA bases are exchanged for their corresponding
bases except in the case of thymine (T), for which RNA substitutes
uracil (U).[92] In eukaryotes, a large part of DNA (e.g., >98% in
humans) contain non-coding called introns, which do not serve as
patterns for protein sequences. The coding regions or exons are
interspersed along with the introns in the primary transcript (or pre-
mRNA).[91] Before translation, the pre-mRNA undergoes further
processing whereby the introns are removed (or spliced out), leaving
only the spliced exons in the mature mRNA strand.[91]

The translation of mRNA to protein occurs in ribosomes, whereby the


transcribed mRNA strand specifies the sequence of amino acids
:
within proteins using the genetic code. Gene products are often
proteins, but in non-protein-coding genes such as transfer RNA
(tRNA) and small nuclear RNA (snRNA), the product is a functional
non-coding RNA.[93][94]

Gene regulation

The regulation of gene expression (or


gene regulation) by environmental factors
and during different stages of
development can occur at each step of the
process such as transcription, RNA
splicing, translation, and post-translational
modification of a protein.[95] Regulation of various stages of
gene expression
The ability of gene transcription to be
regulated allows for the conservation of energy as cells will only
make proteins when needed.[95] Gene expression can be influenced
by positive or negative regulation, depending on which of the two
types of regulatory proteins called transcription factors bind to the
DNA sequence close to or at a promoter.[95] A cluster of genes that
share the same promoter is called an operon, found mainly in
prokaryotes and some lower eukaryotes (e.g., Caenorhabditis
elegans).[95][96] It was first identified in Escherichia coli—a
prokaryotic cell that can be found in the intestines of humans and
other animals—in the 1960s by François Jacob and Jacques Monod.
[95] They studied the prokaryotic cell's lac operon, which is part of

three genes (lacZ, lacY, and lacA) that encode three lactose-
metabolizing enzymes (β-galactosidase, β-galactoside permease,
and β-galactoside transacetylase).[95] In positive regulation of gene
expression, the activator is the transcription factor that stimulates
transcription when it binds to the sequence near or at the promoter.
In contrast, negative regulation occurs when another transcription
:
factor called a repressor binds to a DNA sequence called an
operator, which is part of an operon, to prevent transcription. When a
repressor binds to a repressible operon (e.g., trp operon), it does so
only in the presence of a corepressor. Repressors can be inhibited by
compounds called inducers (e.g., allolactose), which exert their
effects by binding to a repressor to prevent it from binding to an
operator, thereby allowing transcription to occur.[95] Specific genes
that can be activated by inducers are called inducible genes (e.g.,
lacZ or lacA in E. coli), which are in contrast to constitutive genes
that are almost always active.[95] In contrast to both, structural
genes encode proteins that are not involved in gene regulation.[95]

In prokaryotic cells, transcription is regulated by proteins called


sigma factors, which bind to RNA polymerase and direct it to specific
promoters.[95] Similarly, transcription factors in eukaryotic cells can
also coordinate the expression of a group of genes, even if the genes
themselves are located on different chromosomes.[95] Coordination
of these genes can occur as long as they share the same regulatory
DNA sequence that bind to the same transcription factors.[95]
Promoters in eukaryotic cells are more diverse but tend to contain a
core sequence that RNA polymerase can bind to, with the most
common sequence being the TATA box, which contains multiple
repeating A and T bases.[95] Specifically, RNA polymerase II is the
RNA polymerase that binds to a promoter to initiate transcription of
protein-coding genes in eukaryotes, but only in the presence of
multiple general transcription factors, which are distinct from the
transcription factors that have regulatory effects, i.e., activators and
repressors.[95] In eukaryotic cells, DNA sequences that bind with
activators are called enhances whereas those sequences that bind
with repressors are called silencers.[95] Transcription factors such as
nuclear factor of activated T-cells (NFAT) are able to identify specific
nucleotide sequence based on the base sequence (e.g.,
:
CGAGGAAAATTG for NFAT) of the binding site, which determines
the arrangement of the chemical groups within that sequence that
allows for specific DNA-protein interactions.[95] The expression of
transcription factors is what underlies cellular differentiation in a
developing embryo.[95]

In addition to regulatory events involving the promoter, gene


expression can also be regulated by epigenetic changes to
chromatin, which is a complex of DNA and protein found in
eukaryotic cells.[95]

Post-transcriptional control of mRNA can involve the alternative


splicing of primary mRNA transcripts, resulting in a single gene
giving rise to different mature mRNAs that encode a family of
different proteins.[95][97] A well-studied example is the Sxl gene in
Drosophila, which determines the sex in these animals. The gene
itself contains four exons and alternative splicing of its pre-mRNA
transcript can generate two active forms of the Sxl protein in female
flies and one in inactive form of the protein in males.[95] Another
example is the human immunodeficiency virus (HIV), which has a
single pre-mRNA transcript that can generate up to nine proteins as
a result of alternative splicing.[95] In humans, eighty percent of all
21,000 genes are alternatively spliced.[95] Given that both
chimpanzees and humans have a similar number of genes, it is
thought that alternative splicing might have contributed to the
latter's complexity due to the greater number of alternative splicing
in the human brain than in the brain of chimpanzees.[95]

Translation can be regulated in three known ways, one of which


involves the binding of tiny RNA molecules called microRNA (miRNA)
to a target mRNA transcript, which inhibits its translation and causes
it to degrade.[95] Translation can also be inhibited by the
modification of the 5’ cap by substituting the modified guanosine
:
triphosphate (GTP) at the 5’ end of an mRNA for an unmodified GTP
molecule.[95] Finally, translational repressor proteins can bind to
mRNAs and prevent them from attaching to a ribosome, thereby
blocking translation.[95]

Once translated, the stability of proteins can be regulated by being


targeted for degradation.[95] A common example is when an enzyme
attaches a regulatory protein called ubiquitin to the lysine residue of
a targeted protein.[95] Other ubiquitins then attached to the primary
ubiquitin to form a polyubiquitinated protein, which then enters a
much larger protein complex called proteasome.[95] Once the
polyubiquitinated protein enters the proteasome, the polyubiquitin
detaches from the target protein, which is unfolded by the
proteasome in an ATP-dependent manner, allowing it to be
hydrolyzed by three proteases.[95]

Genomes

Further information: Genomics

A genome is an organism's complete set


of DNA, including all of its genes.[98]
Sequencing and analysis of genomes can
be done using high throughput DNA
sequencing and bioinformatics to
assemble and analyze the function and
structure of entire genomes.[99][100][101] Composition of the human
The genomes of prokaryotes are small, genome

compact, and diverse. In contrast, the


genomes of eukaryotes are larger and more complex such as having
more regulatory sequences and much of its genome are made up of
non-coding DNA sequences for functional RNA (rRNA, tRNA, and
mRNA) or regulatory sequences. The genomes of various model
:
organisms such as arabidopsis, fruit fly, mice, nematodes, and yeast
have been sequenced. The Human Genome Project was a major
undertaking by the international scientific community to sequence
the entire human genome, which was completed in 2003.[102] The
sequencing of the human genome has yielded practical applications
such as DNA fingerprinting, which can be used for paternity testing
and forensics. In medicine, sequencing of the entire human genome
has allowed for the identification of mutations that cause tumors as
well as genes that cause a specific genetic disorder.[102] The
sequencing of genomes from various organisms has led to the
emergence of comparative genomics, which aims to draw
comparisons of genes from the genomes of those different
organisms.[102]

Many genes encode more than one protein, with posttranslational


modifications increasing the diversity of proteins within a cell. An
organism's proteome is its entire set of proteins expressed by its
genome and proteomics seeks to study the complete set of proteins
produced by an organism.[102] Because many proteins are enzymes,
their activities tend to affects the concentrations of substrates and
products. Thus, as the proteome changes, so do the amount of small
molecules or metabolites.[102] The complete set of small molecules
in a cell or organism is called a metabolome and metabolomics is the
study of the metabolome in relation to the physiological activity of a
cell or organism.[102]

Biotechnology

Further information: Molecular biology

Biotechnology is the use of cells or organisms to develop products


for humans.[103] One commonly used technology with wide
applications is the creation of recombinant DNA, which is a DNA
:
molecule assembled from two or more
sources in a laboratory. Before the advent
of polymerase chain reaction, biologists
would manipulate DNA by cutting it into
smaller fragments using restriction
enzymes. They would then purify and
analyze the fragments using gel
electrophoresis and then later recombine Construction of recombinant
DNA, in which a foreign DNA
the fragments into a novel DNA sequence fragment is inserted into a
using DNA ligase.[103] The recombinant plasmid vector

DNA is then cloned by inserting it into a host cell, a process known


as transformation if the host cells were bacteria such as E. coli, or
transfection if the host cells were eukaryotic cells like yeast, plant, or
animal cells. Once the host cell or organism has received and
integrated the recombinant DNA, it is described as transgenic.[103]

A recombinant DNA can be inserted in one of two ways. A common


method is to simply insert the DNA into a host chromosome, with the
site of insertion being random.[103] Another approach would be to
insert the recombinant DNA as part of another DNA sequence called
a vector, which then integrates into the host chromosome or has its
own origin of DNA replication, thereby allowing to replicate
independently of the host chromosome.[103] Plasmids from bacterial
cells such as E. coli are typically used as vectors due to their
relatively small size (e.g. 2000-6000 base pairs in E. coli), presence
of restriction enzymes, genes that are resistant to antibiotics, and
the presence of an origin of replication.[103] A gene coding for a
selectable marker such as antibiotic resistance is also incorporated
into the vector.[103] Inclusion of this market allows for the selection of
only those host cells that contained the recombinant DNA while
discarding those that do not.[103] Moreover, the marker also serves
as a reporter gene that once expressed, can be easily detected and
:
measured.[103]

Once the recombinant DNA is inside individual bacterial cells, those


cells are then plated and allowed to grow into a colony that contains
millions of transgenic cells that carry the same recombinant DNA.
[104] These transgenic cells then produce large quantities of the

transgene product such as human insulin, which was the first


medicine to be made using recombinant DNA technology.[103]

One of the goals of molecular cloning is to identify the function of


specific DNA sequences and the proteins they encode.[103] For a
specific DNA sequence to be studied and manipulated, millions of
copies of DNA fragments containing that DNA sequence need to be
made.[103] This involves breaking down an intact genome, which is
much too large to be introduced into a host cell, into smaller DNA
fragments. Although no longer intact, the collection of these DNA
fragments still make up an organism's genome, with the collection
itself being referred to as a genomic library, due to the ability to
search and retrieve specific DNA fragments for further study,
analogous to the process of retrieving a book from a regular library.
[103] DNA fragments can be obtained using restriction enzymes and

other processes such as mechanical shearing. Each obtained


fragment is then inserted into a vector that is taken up by a bacterial
host cell. The host cell is then allowed to proliferate on a selective
medium (e.g., antibiotic resistance), which produces a colony of
these recombinant cells, each of which contains many copies of the
same DNA fragment.[103] These colonies can be grown by spreading
them over a solid medium in Petri dishes, which are incubated at a
suitable temperature. One dish alone can hold thousands of bacterial
colonies, which can be easily screened for a specific DNA sequence.
[103] The sequence can be identified by first duplicating a Petri dish

with bacterial colonies and then exposing the DNA of the duplicated
colonies for hybridization, which involves labeling them with
:
complementary radioactive or fluorescent nucleotides.[103]

Smaller DNA libraries that contain genes from a specific tissue can
be created using complementary DNA (cDNA).[103] The collection of
these cDNAs from a specific tissue at a particular time is called a
cDNA library, which provides a "snapshot" of transcription patterns
of cells at a specific location and time.[103]

Other biotechnology tools include DNA microarrays, expression


vectors, synthetic genomics, and CRISPR gene editing.[103][105]
Other approaches such as pharming can produce large quantities of
medically useful products through the use of genetically modified
organisms.[103] Many of these other tools also have wide
applications such as creating medically useful proteins, or improving
plant cultivation and animal husbandry.[103]

Genes, development, and evolution

Further information: Evolutionary developmental biology

Development is the process by which a


multicellular organism (plant or animal) goes
through a series of a changes, starting from
a single cell, and taking on various forms
that are characteristic of its life cycle.[107]
Model of concentration
There are four key processes that underlie gradient building up; fine
development: Determination, differentiation, yellow-orange outlines are cell
[106]
morphogenesis, and growth. Determination boundaries.
sets the developmental fate of a cell, which becomes more
restrictive during development. Differentiation is the process by
which specialized cells from less specialized cells such as stem cells.
[108][109] Stem cells are undifferentiated or partially differentiated

cells that can differentiate into various types of cells and proliferate
indefinitely to produce more of the same stem cell.[110] Cellular
:
differentiation dramatically changes a cell's size, shape, membrane
potential, metabolic activity, and responsiveness to signals, which
are largely due to highly controlled modifications in gene expression
and epigenetics. With a few exceptions, cellular differentiation
almost never involves a change in the DNA sequence itself.[111] Thus,
different cells can have very different physical characteristics despite
having the same genome. Morphogenesis, or development of body
form, is the result of spatial differences in gene expression.[107]
Specially, the organization of differentiated tissues into specific
structures such as arms or wings, which is known as pattern
formation, is governed by morphogens, signaling molecules that
move from one group of cells to surrounding cells, creating a
morphogen gradient as described by the French flag model.
Apoptosis, or programmed cell death, also occurs during
morphogenesis, such as the death of cells between digits in human
embryonic development, which frees up individual fingers and toes.
Expression of transcription factor genes can determine organ
placement in a plant and a cascade of transcription factors
themselves can establish body segmentation in a fruit fly.[107]

A small fraction of the genes in an organism's genome called the


developmental-genetic toolkit control the development of that
organism. These toolkit genes are highly conserved among phyla,
meaning that they are ancient and very similar in widely separated
groups of animals. Differences in deployment of toolkit genes affect
the body plan and the number, identity, and pattern of body parts.
Among the most important toolkit genes are the Hox genes. Hox
genes determine where repeating parts, such as the many vertebrae
of snakes, will grow in a developing embryo or larva.[112] Variations in
the toolkit may have produced a large part of the morphological
evolution of animals. The toolkit can drive evolution in two ways. A
toolkit gene can be expressed in a different pattern, as when the
:
beak of Darwin's large ground-finch was enlarged by the BMP gene,
[113] or when snakes lost their legs as Distal-less (Dlx) genes became

under-expressed or not expressed at all in the places where other


reptiles continued to form their limbs.[114] Or, a toolkit gene can
acquire a new function, as seen in the many functions of that same
gene, distal-less, which controls such diverse structures as the
mandible in vertebrates,[115][116] legs and antennae in the fruit fly,[117]
and eyespot pattern in butterfly wings.[118] Given that small changes
in toolbox genes can cause significant changes in body structures,
they have often enabled convergent or parallel evolution.

Evolution

Evolutionary processes

Further information: Evolutionary biology

A central organizing concept in biology is


that life changes and develops through
evolution, which is the change in heritable
characteristics of populations over
successive generations.[119][120] Evolution
is now used to explain the great variations
of life on Earth. The term evolution was
introduced into the scientific lexicon by Natural selection for darker traits

Jean-Baptiste de Lamarck in 1809.[121][122] He proposed that


evolution occurred as a result of inheritance of acquired
characteristics, which was unconvincing but there were no
alternative explanations at the time.[121] Charles Darwin, an English
naturalist, had returned to England in 1836 from his five-year travels
on the HMS Beagle where he studied rocks and collected plants and
animals from various parts of the world such as the Galápagos
Islands.[121] He had also read Principles of Geology by Charles Lyell
:
and An Essay on the Principle of Population by Thomas Malthus and
was influenced by them.[123] Based on his observations and
readings, Darwin began to formulate his theory of evolution by
natural selection to explain the diversity of plants and animals in
different parts of the world.[121][123] Alfred Russel Wallace, another
English naturalist who had studied plants and animals in the Malay
Archipelago, also came to the same idea, but later and
independently of Darwin.[121] Both Darwin and Wallace jointly
presented their essay and manuscript, respectively, at the Linnaean
Society of London in 1858, giving them both credit for their
discovery of evolution by natural selection.[121][124][125][126][127]
Darwin would later publish his book On the Origin of Species in 1859,
which explained in detail how the process of evolution by natural
selection works.[121]

To explain natural selection, Darwin drew an analogy with humans


modifying animals through artificial selection, whereby animals were
selectively bred for specific traits, which has given rise to individuals
that no longer resemble their wild ancestors.[123] Darwin argued that
in the natural world, it was nature that played the role of humans in
selecting for specific traits. He came to this conclusion based on two
observations and two inferences.[123] First, members of any
population tend to vary with respect to their heritable traits. Second,
all species tend to produce more offspring than can be supported by
their respective environments, resulting in many individuals not
surviving and reproducing.[123] Based on these observations, Darwin
inferred that those individuals who possessed heritable traits that are
better adapted to their environments are more likely to survive and
produce more offspring than other individuals.[123] He further
inferred that the unequal or differential survival and reproduction of
certain individuals over others will lead to the accumulation of
favorable traits over successive generations, thereby increasing the
:
match between the organisms and their environment.[123][128][129]
Thus, taken together, natural selection is the differential survival and
reproduction of individuals in subsequent generations due to
differences in or more heritable traits.[130][123][121]

Darwin was not aware of Mendel's work of inheritance and so the


exact mechanism of inheritance that underlie natural selection was
not well-understood[131] until the early 20th century when the
modern synthesis reconciled Darwinian evolution with classical
genetics, which established a neo-Darwinian perspective of
evolution by natural selection.[130] This perspective holds that
evolution occurs when there are changes in the allele frequencies
within a population of interbreeding organisms. In the absence of any
evolutionary process acting on a large random mating population,
the allele frequencies will remain constant across generations as
described by the Hardy–Weinberg principle.[132]

Another process that drives evolution is genetic drift, which is the


random fluctuations of allele frequencies within a population from
one generation to the next.[133] When selective forces are absent or
relatively weak, allele frequencies are equally likely to drift upward or
downward at each successive generation because the alleles are
subject to sampling error.[134] This drift halts when an allele
eventually becomes fixed, either by disappearing from the
population or replacing the other alleles entirely. Genetic drift may
therefore eliminate some alleles from a population due to chance
alone.

Speciation

A species is a group of organisms that mate with one another and


speciation is the process by which one lineage splits into two
lineages as a result of having evolved independently from each other.
:
[135]For speciation to occur,
there has to be reproductive
isolation.[135] Reproductive
isolation can result from
incompatibilities between genes
as described by Bateson–
Dobzhansky–Muller model.
Reproductive isolation also tends
to increase with genetic
divergence. Speciation can occur
when there are physical barriers Comparison of allopatric, peripatric,
that divide an ancestral species, parapatric and sympatric speciation
a process known as allopatric speciation.[135] In contrast, sympatric
speciation occurs in the absence of physical barriers.

Pre-zygotic isolation such as mechanical, temporal, behavioral,


habitat, and gametic isolations can prevent different species from
hybridizing.[135] Similarly, post-zygotic isolations can result in
hybridization being selected against due to the lower viability of
hybrids or hybrid infertility (e.g., mule). Hybrid zones can emerge if
there were to be incomplete reproductive isolation between two
closely related species.

Phylogeny

Further information: Phylogenetics and Biodiversity


:
Phylogenetic tree showing the domains of bacteria, archaea, and
eukaryotes

A phylogeny is an evolutionary history of a specific group of


organisms or their genes.[136] It can be represented using a
phylogenetic tree, which is a diagram showing lines of descent
among organisms or their genes. Each line drawn on the time axis of
a tree represents a lineage of descendants of a particular species or
population. When a lineage divides into two, it is represented as a
node (or split) on the phylogenetic tree. The more splits there are
over time, the more branches there will be on the tree, with the
common ancestor of all the organisms in that tree being represented
by the root of that tree. Phylogenetic trees may portray the
evolutionary history of all life forms, a major evolutionary group (e.g.,
insects), or an even smaller group of closely related species. Within a
tree, any group of species designated by a name is a taxon (e.g.,
humans, primates, mammals, or vertebrates) and a taxon that
consists of all its evolutionary descendants is a clade, otherwise
known as a monophyletic taxon.[136] Closely related species are
referred to as sister species and closely related clades are sister
clades. In contrast to a monophyletic group, a polyphyletic group
does not include its common ancestor whereas a paraphyletic group
does not include all the descendants of a common ancestor.[136]
:
Phylogenetic trees are the basis for comparing and grouping
different species.[136] Different species that share a feature inherited
from a common ancestor are described as having homologous
features (or synapomorphy).[137][138][136] Homologous features may
be any heritable traits such as DNA sequence, protein structures,
anatomical features, and behavior patterns. A vertebral column is an
example of a homologous feature shared by all vertebrate animals.
Traits that have a similar form or function but were not derived from a
common ancestor are described as analogous features. Phylogenies
can be reconstructed for a group of organisms of primary interests,
which are called the ingroup. A species or group that is closely
related to the ingroup but is phylogenetically outside of it is called
the outgroup, which serves a reference point in the tree. The root of
the tree is located between the ingroup and the outgroup.[136] When
phylogenetic trees are reconstructed, multiple trees with different
evolutionary histories can be generated. Based on the principle of
Parsimony (or Occam's razor), the tree that is favored is the one with
the fewest evolutionary changes needed to be assumed over all
traits in all groups. Computational algorithms can be used to
determine how a tree might have evolved given the evidence.[136]

Phylogeny provides the basis of biological classification, which is


based on Linnaean taxonomy that was developed by Carl Linnaeus in
the 18th century.[136] This classification system is rank-based, with
the highest rank being the domain followed by kingdom, phylum,
class, order, family, genus, and species.[136] All organisms can be
classified as belonging to one of three domains: Archaea (originally
Archaebacteria); bacteria (originally eubacteria), or eukarya (includes
the protist, fungi, plant, and animal kingdoms).[139] A binomial
nomenclature is used to classify different species. Based on this
system, each species is given two names, one for its genus and
another for its species.[136] For example, humans are Homo sapiens,
:
with Homo being the genus and sapiens being the species. By
convention, the scientific names of organisms are italicized, with only
the first letter of the genus capitalized.[140][141]

History of life

Life timeline

This box:
view
talk
edit

Water

← Earth formed
Single-celled life

← Earliest water
Photosynthesis

Earliest
Eukaryotes ←
known life

Multicellular life
LHB

meteorites

P
l Earliest
−4500 — ←
a oxygen
– n
— t
s Pongola
– ←
glaciation*
:
−4000 — Arthropods Molluscs
Atmospheric
– Flowers ←
oxygen
— Dinosaurs
– Huronian

−3500 — Mammal glaciation*

– s
Birds Sexual
— ←
reproduction
– Primates

−3000 — H
a Earliest
– d ← multicellular
e life

a

n
−2500 — ← Earliest fungi


— Earliest

A plants

r
−2000 — c
h Earliest
– ←
animals
e

a
– n Cryogenian

−1500 — ice age*


— ←
Ediacaran
biota

−1000 —
Cambrian
– P ←
explosion
— r
o

:
−500 — t ← Andean
glaciation*
– e
r
— o
Earliest
– z ←
tetrapods
o
0—
i
c Karoo ice

age*
P
h
a
Earliest apes
n ←
/ humans
e
r
o Quaternary

z ice age*
o
i
c

(million years ago)


*Ice Ages

The history of life on Earth traces the processes by which organisms


have evolved from the earliest emergence of life to present day.
Earth formed about 4.5 billion years ago and all life on Earth, both
living and extinct, descended from a last universal common ancestor
that lived about 3.5 billion years ago.[142][143] The dating of the
Earth's history can be done using several geological methods such
as stratigraphy, radiometric dating, and paleomagnetic dating.[144]
:
Based on these methods, geologists have developed a geologic time
scale that divides the history of the Earth into major divisions,
starting with four eons (Hadean, Archean, Proterozoic, and
Phanerozoic), the first three of which are collectively known as the
Precambrian, which lasted approximately 4 billion years.[144] Each
eon can be divided into eras, with the Phanerozoic eon that began
542 million years ago being subdivided into Paleozoic, Mesozoic,
and Cenozoic eras.[144] These three eras together comprise eleven
periods (Cambrian, Ordovician, Silurian, Devonian, Carboniferous,
Permian, Triassic, Jurassic, Cretaceous, Tertiary, and Quaternary)
and each period into epochs.[144]

The similarities among all known present-day species indicate that


they have diverged through the process of evolution from their
common ancestor.[145] Biologists regard the ubiquity of the genetic
code as evidence of universal common descent for all bacteria,
archaea, and eukaryotes.[146][10][147][148] Microbal mats of coexisting
bacteria and archaea were the dominant form of life in the early
Archean epoch and many of the major steps in early evolution are
thought to have taken place in this environment.[149] The earliest
evidence of eukaryotes dates from 1.85 billion years ago,[150][151] and
while they may have been present earlier, their diversification
accelerated when they started using oxygen in their metabolism.
Later, around 1.7 billion years ago, multicellular organisms began to
appear, with differentiated cells performing specialised functions.
[152]

Algae-like multicellular land plants are dated back even to about 1


billion years ago,[153] although evidence suggests that
microorganisms formed the earliest terrestrial ecosystems, at least
2.7 billion years ago.[154] Microorganisms are thought to have paved
the way for the inception of land plants in the Ordovician period.
Land plants were so successful that they are thought to have
:
contributed to the Late Devonian extinction event.[155]

Ediacara biota appear during the Ediacaran period,[156] while


vertebrates, along with most other modern phyla originated about
525 million years ago during the Cambrian explosion.[157] During the
Permian period, synapsids, including the ancestors of mammals,
dominated the land,[158] but most of this group became extinct in the
Permian–Triassic extinction event 252 million years ago.[159] During
the recovery from this catastrophe, archosaurs became the most
abundant land vertebrates;[160] one archosaur group, the dinosaurs,
dominated the Jurassic and Cretaceous periods.[161] After the
Cretaceous–Paleogene extinction event 66 million years ago killed
off the non-avian dinosaurs,[162] mammals increased rapidly in size
and diversity.[163] Such mass extinctions may have accelerated
evolution by providing opportunities for new groups of organisms to
diversify.[164]

Diversity

Bacteria and Archaea

Further information: Microbiology

Bacteria are a type of cell that constitute a


large domain of prokaryotic
microorganisms. Typically a few
micrometers in length, bacteria have a
number of shapes, ranging from spheres
Bacteria – Gemmatimonas
to rods and spirals. Bacteria were among
aurantiaca (-=1 Micrometer) the first life forms to appear on Earth, and
are present in most of its habitats. Bacteria
inhabit soil, water, acidic hot springs, radioactive waste,[165] and the
deep biosphere of the earth's crust. Bacteria also live in symbiotic
:
and parasitic relationships with plants and animals. Most bacteria
have not been characterised, and only about 27 percent of the
bacterial phyla have species that can be grown in the laboratory.[166]

Archaea constitute the other domain of


prokaryotic cells and were initially
classified as bacteria, receiving the name
archaebacteria (in the Archaebacteria
kingdom), a term that has fallen out of use.
[167] Archaeal cells have unique properties

separating them from the other two


Archaea – Halobacteria
domains, Bacteria and Eukaryota. Archaea
are further divided into multiple
recognized phyla. Archaea and bacteria are generally similar in size
and shape, although a few archaea have very different shapes, such
as the flat and square cells of Haloquadratum walsbyi.[168] Despite
this morphological similarity to bacteria, archaea possess genes and
several metabolic pathways that are more closely related to those of
eukaryotes, notably for the enzymes involved in transcription and
translation. Other aspects of archaeal biochemistry are unique, such
as their reliance on ether lipids in their cell membranes,[169] including
archaeols. Archaea use more energy sources than eukaryotes: these
range from organic compounds, such as sugars, to ammonia, metal
ions or even hydrogen gas. Salt-tolerant archaea (the Haloarchaea)
use sunlight as an energy source, and other species of archaea fix
carbon, but unlike plants and cyanobacteria, no known species of
archaea does both. Archaea reproduce asexually by binary fission,
fragmentation, or budding; unlike bacteria, no known species of
Archaea form endospores.

The first observed archaea were extremophiles, living in extreme


environments, such as hot springs and salt lakes with no other
organisms. Improved molecular detection tools led to the discovery
:
of archaea in almost every habitat, including soil, oceans, and
marshlands. Archaea are particularly numerous in the oceans, and
the archaea in plankton may be one of the most abundant groups of
organisms on the planet.

Archaea are a major part of Earth's life. They are part of the
microbiota of all organisms. In the human microbiome, they are
important in the gut, mouth, and on the skin.[170] Their
morphological, metabolic, and geographical diversity permits them
to play multiple ecological roles: carbon fixation; nitrogen cycling;
organic compound turnover; and maintaining microbial symbiotic
and syntrophic communities, for example.[171]

Protists

Further information: Protistology

Eukaryotes are hypothesized to have split


from archaea, which was followed by their
endosymbioses with bacteria (or
symbiogenesis) that gave rise to
mitochondria and chloroplasts, both of
which are now part of modern day
eukaryotic cells.[172] The major lineages of
eukaryotes diversified in the Precambrian
about 1.5 billion years ago and can be
classified into eight major clades:
alveolates, excavates, stramenopiles, Diversity of protists

plants, rhizarians, amoebozoans, fungi, and animals.[172] Five of


these clades are collectively known as protists, which are mostly
microscopic eukaryotic organisms that are not plants, fungi, or
animals.[172] While it is likely that protists share a common ancestor
(the last eukaryotic common ancestor),[173] protists by themselves
:
do not constitute a separate clade as some protists may be more
closely related to plants, fungi, or animals than they are to other
protists. Like groupings such as algae, invertebrates, or protozoans,
the protist grouping is not a formal taxonomic group but is used for
convenience.[172][174] Most protists are unicellular, which are also
known as microbial eukaryotes.[172]

The alveolates are mostly photosynthetic unicellular protists that


possess sacs called alveoli (hence their name alveolates) that are
located beneath their cell membrane, providing support for the cell
surface.[172] Alveolates comprise several groups such as
dinoflagellates, apicomplexans, and ciliates. Dinoflagellates are
photosynthetic and can be found in the ocean where they play a role
as primary producers of organic matter.[172] Apicomplexans are
parasitic alveolates that possess an apical complex, which is a group
of organelles located in the apical end of the cell.[172] This complex
allows apicomplexans to invade their hosts' tissues. Ciliates are
alveolates that possess numerous hair-like structure called cilia. A
defining characteristic of ciliates is the presence of two types of
nuclei in each ciliate cell. A commonly studied ciliate is the
paramecium.[172]

The excavates are groups of protists that began to diversify


approximately 1.5 billion years ago shortly after the origin of the
eukaryotes.[172] Some excavates do not possess mitochondria,
which are thought to have been lost over the course of evolution as
these protists still possess nuclear genes that are associated with
mitochondria.[172] The excavates comprise several groups such as
diplomonads, parabasalids, heteroloboseans, euglenids, and
kinetoplastids.[172]

Stramenopiles, most of which can be characterized by the presence


of tubular hairs on the longer of their two flagella, include diatoms
:
and brown algae.[172] Diatoms are primary producers and contribute
about one-fifth of all photosynthetic carbon fixation, making them a
major component of phytoplankton.[172]

Rhizarians are mostly unicellular and aquatic protists that typically


contain long, thin pseudopods.[172] The rhizarians comprise three
main groups: cercozoans, foraminiferans, and radiolarians.[172]

Amoebozoans are protists with a body form characterized by the


presence lobe-shaped pseudopods, which help them to move.[172]
They include groups such as loboseans and slime molds (e.g.,
plasmodial slime mold and cellular slime molds).[172]

Plant diversity

Further information: Botany

Plants are mainly multicellular organisms,


predominantly photosynthetic eukaryotes
of the kingdom Plantae, which would
exclude fungi and some algae. A shared
derived trait (or synapomorphy) of Plantae
is the primary endosymbiosis of a
cyanobacterium into an early eukaryote
about one billion years ago, which gave
rise to chloroplasts.[175] The first several
clades that emerged following primary
endosymbiosis were aquatic and most of
the aquatic photosynthetic eukaryotic Diversity of plants
organisms are collectively described as
algae, which is a term of convenience as not all algae are closely
related.[175] Algae comprise several distinct clades such as
glaucophytes, which are microscopic freshwater algae that may have
resembled in form to the early unicellular ancestor of Plantae.[175]
:
Unlike glaucophytes, the other algal clades such as red and green
algae are multicellular. Green algae comprise three major clades:
chlorophytes, coleochaetophytes, and stoneworts.[175]

Land plants (embryophytes) first appeared in terrestrial


environments approximately 450 to 500 million years ago.[175] A
synapomorphy of land plants is an embryo that develops under the
protection of tissues of its parent plant.[175] Land plants comprise ten
major clades, seven of which constitute a single clade known as
vascular plants (or tracheophytes) as they all have tracheids, which
are fluid-conducting cells, and a well-developed system that
transports materials throughout their bodies.[175] In contrast, the
other three clades are nonvascular plants as they do not have
tracheids.[175] They also do not constitute a single clade.[175]

Nonvascular plants include liverworts, mosses, and hornworts. They


tend to be found in areas where water is readily available.[175] Most
live on soil or even on vascular plants themselves. Some can grow on
bare rock, tree trunks that are dead or have fallen, and even
buildings.[175] Most nonvascular plants are terrestrial, with a few
living in freshwater environments and none living in the oceans.[175]

The seven clades (or divisions) that make up vascular plants include
horsetails and ferns, which together can be grouped as a single
clade called monilophytes.[175] Seed plants (or spermatophyte)
comprise the other five divisions, four of which are grouped as
gymnosperms and one is angiosperms. Gymnosperms includes
conifers, cycads, Ginkgo, and gnetophytes. Gymnosperm seeds
develop either on the surface of scales or leaves, which are often
modified to form cones, or solitary as in yew, Torreya, Ginkgo.[176]
Angiosperms are the most diverse group of land plants, with 64
orders, 416 families, approximately 13,000 known genera and
300,000 known species.[177] Like gymnosperms, angiosperms are
:
seed-producing plants. They are distinguished from gymnosperms
by having characteristics such as flowers, endosperm within their
seeds, and production of fruits that contain the seeds.

Fungi

Further information: Mycology

Fungi are eukaryotic organisms that digest


foods outside of their bodies.[178] They do
so through a process called absorptive
heterotrophy whereby they would first
secrete digestive enzymes that break
down large food molecules before
absorbing them through their cell
Diversity of fungi. Clockwise from
membranes. Many fungi are also saprobes top left: Amanita muscaria, a
as they are able to take in nutrients from basidiomycete; Sarcoscypha
coccinea, an ascomycete; bread
dead organic matter and are hence, the covered in mold; chytrid;
principal decomposers in ecological Aspergillus conidiophore.
systems. [178] Some fungi are parasites by
absorbing nutrients from living hosts while others are mutualists.[178]
Fungi, along with two other lineages, choanoflagellates and animals,
can be grouped as opisthokonts. A synapomorphy that distinguishes
fungi from other two opisthokonts is the presence of chitin in their
cell walls.[178]

Most fungi are multicellular but some are unicellular such as yeasts,
which live in liquid or moist environments and are able to absorb
nutrients directly into their cell surfaces.[178] Multicellular fungi, on
the other hand, have a body called mycelium, which is composed of
a mass of individual tubular filaments called hyphae that allows for
nutrient absorption to occur.[178]

Fungi can be divided into six major groups based on their life cycles:
:
microsporidia, chytrids, zygospore fungi (Zygomycota), arbuscular
mycorrhizal fungi (Glomeromycota), sac fungi (Ascomycota), and
club fungi (Basidiomycota).[178] Fungi are classified by the particular
processes of sexual reproduction they use. The usual cellular
products of meiosis during sexual reproduction are spores that are
adapted to survive inclement times and to spread. A principal
adaptive benefit of meiosis during sexual reproduction in the
Ascomycota and Basidiomycota was proposed to be the repair of
DNA damage through meiotic recombination.[179]

The fungus kingdom encompasses an enormous diversity of taxa


with varied ecologies, life cycle strategies, and morphologies ranging
from unicellular aquatic chytrids to large mushrooms. However, little
is known of the true biodiversity of Kingdom Fungi, which has been
estimated at 2.2 million to 3.8 million species.[180] Of these, only
about 148,000 have been described,[181] with over 8,000 species
known to be detrimental to plants and at least 300 that can be
pathogenic to humans.[182]

Animal diversity

Further information: Zoology

Animals are multicellular eukaryotic organisms that form the


kingdom Animalia. With few exceptions, animals consume organic
material, breathe oxygen, are able to move, can reproduce sexually,
and grow from a hollow sphere of cells, the blastula, during
embryonic development. Over 1.5 million living animal species have
been described—of which around 1 million are insects—but it has
been estimated there are over 7 million animal species in total. They
have complex interactions with each other and their environments,
forming intricate food webs.

Animals can be distinguished into two groups based on their


:
developmental characteristics.[183] For
instance, embryos of diploblastic animals
such as ctenophores, placeozoans, and
cnidarians have two cell layers (ectoderm
and endoderm) whereas the embryos of
triploblastic animals have three tissue
layers (ectoderm, mesoderm, and
endoderm), which is a synapomorphy of
these animals.[183] Triploblastic animals
can be further divided into two major
clades based on based on the pattern of
gastrulation, whereby a cavity called a
blastopore is formed from the indentation Diversity of animals. From top to
bottom, first column:
of a blastula. In protostomes, the Echinoderm, cnidaria, bivalve,
blastopore gives rise to the mouth, which tardigrade, crustacean, and
arachnid. Second column:
is then followed by the formation of the
Sponge, insect, mammal,
anus.[183] In deuterostomes, the bryozoa, acanthocephala, and
blastopore gives rise to the anus, followed flatworm. Third column:
Cephalopod, annelid, tunicate,
by the formation of the mouth.[183] fish, bird, and phoronida.

Animals can also be differentiated based on their body plan,


specifically with respect to four key features: symmetry, body cavity,
segmentation, and appendages.[183] The bodies of most animals are
symmetrical, with symmetry being either radial or bilateral.[183]
Triploblastic animals can be divided into three types based on their
body cavity: acoelomate, pseudocoelomate, and coelomate.[183]
Segmentation can be observed in the bodies of many animals, which
allows for specialization of different parts of the body as well as
allowing the animal to change the shape of its body to control its
movements.[183] Finally, animals can be distinguished based on the
type and location of their appendages such as antennae for sensing
the environment or claws for capturing prey.[183]
:
Sponges, the members of the phylum Porifera, are a basal Metazoa
(animal) clade as a sister of the diploblasts.[184][185][186][187][188]
They are multicellular organisms that have bodies full of pores and
channels allowing water to circulate through them, consisting of
jelly-like mesohyl sandwiched between two thin layers of cells.

The majority (~97%) of animal species are invertebrates,[189] which


are animals that do not have a vertebral column (or backbone or
spine), derived from the notochord. This includes all animals apart
from the subphylum Vertebrata. Familiar examples of invertebrates
include sponges, cnidarians (hydras, jellyfishes, sea anemones, and
corals), mollusks (chitons, snail, bivalves, squids, and octopuses),
annelids (earthworms and leeches), and arthropods (insects,
arachnids, crustaceans, and myriapods). Many invertebrate taxa
have a greater number and variety of species than the entire
subphylum of Vertebrata.[190]

In contrast, vertebrates comprise all species of animals within the


subphylum Vertebrata, which are chordates with vertebral columns.
These animals have four key features, which are an anterior skull with
a brain, a rigid internal skeleton supported by a vertebral column that
encloses a spinal cord, internal organs suspended in a coelom, and a
well-developed circulatory system driven by a single large heart.[183]
Vertebrates represent the overwhelming majority of the phylum
Chordata, with currently about 69,963 species described.[191]
Vertebrates comprise different major groups that include jawless
fishes (not including hagfishes), jawed vertebrates such as
cartilaginous fishes (sharks, rays, and ratfish), bony fishes, tetrapods
such as amphibians, reptiles, birds, and mammals.[183]

The two remaining groups of jawless fishes that have survived


beyond the Devonian period are hagfishes and lamprey, which are
collectively known as cyclostomes (for circled mouths).[183] Both
:
groups of animals have elongated eel-like bodies with no paired fins.
[183] However, because hagfishes have a weak circulatory system

with three accessory hearts, a partial skull with no cerebellum, no


jaws or stomach, and no jointed vertebrae, some biologists do not
classify them as vertebrates but instead as a sister group of
vertebrates.[183] In contrast, lampreys have a complete skull and a
distinct vertebrae that is cartilaginous.[183]

Mammals have four key features that distinguish them from other
animals such as sweat glands, mammary glands, hair, and a four-
chambered heart.[183] Small and medium-sized mammals used to
co-exist with large dinosaurs in much of the Mesozoic era but soon
radiated following the mass extinction of dinosaurs at the end of the
Cretaceous period.[183] There are approximately 57,000 mammal
species, which can be divided into two primary groups: prototherians
and therians. Prototherians do not possess nipples on their
mammary but instead secrete milk onto their skin, allowing their
offspring to lap if off their furs.[183] They also lack a placenta, lays
eggs, and have sprawling legs. Currently, there only five known
species of prototherians (platypus and four species of echidnas).
[183] The therian clade is viviparous and can be further divided into

two groups: marsupials and eutherians.[183] Marsupial females have


a ventral pouch to carry and feed their offspring. Eutherians form the
majority of mammals and include major groups such as rodents,
bats, even-toed ungulates and cetaceans, shrews and moles,
primates, carnivores, rabbits, African insectivores, spiny insectivores,
armadillos, treeshrews, odd-toed ungulates, long-nosed
insectivores, anteaters and sloths, pangolins, hyraxes, sirenians,
elephants, colugos, and aardvark.[183]

Viruses

Further information: Virology


:
Viruses are submicroscopic infectious
agents that replicate inside the cells of
organisms.[192] Viruses infect all types of
life forms, from animals and plants to
microorganisms, including bacteria and
archaea.[193][194] More than 6,000 virus
species have been described in detail.[195]
Viruses are found in almost every
ecosystem on Earth and are the most
numerous type of biological entity.[196][197]
Bacteriophages attached to a
bacterial cell wall
When infected, a host cell is forced to
rapidly produce thousands of identical copies of the original virus.
When not inside an infected cell or in the process of infecting a cell,
viruses exist in the form of independent particles, or virions,
consisting of the genetic material (DNA or RNA), a protein coat
called capsid, and in some cases an outside envelope of lipids. The
shapes of these virus particles range from simple helical and
icosahedral forms to more complex structures. Most virus species
have virions too small to be seen with an optical microscope, as they
are one-hundredth the size of most bacteria.

The origins of viruses in the evolutionary history of life are unclear:


some may have evolved from plasmids—pieces of DNA that can
move between cells—while others may have evolved from bacteria.
In evolution, viruses are an important means of horizontal gene
transfer, which increases genetic diversity in a way analogous to
sexual reproduction.[198] Because viruses possess some but not all
characteristics of life, they have been described as "organisms at the
edge of life",[199] and as self-replicators.[200]

Viruses can spread in many ways. One transmission pathway is


through disease-bearing organisms known as vectors: for example,
:
viruses are often transmitted from plant to plant by insects that feed
on plant sap, such as aphids; and viruses in animals can be carried
by blood-sucking insects. Influenza viruses are spread by coughing
and sneezing. Norovirus and rotavirus, common causes of viral
gastroenteritis, are transmitted by the faecal–oral route, passed by
hand-to-mouth contact or in food or water. Viral infections in animals
provoke an immune response that usually eliminates the infecting
virus. Immune responses can also be produced by vaccines, which
confer an artificially acquired immunity to the specific viral infection.

Plant form and function

Plant body

Further information: Plant morphology, Plant anatomy, and Plant


physiology

The plant body is made up of organs that


can be organized into two major organ
systems: a root system and a shoot
system.[201] The root system anchors the
plants into place. The roots themselves
absorb water and minerals and store
photosynthetic products. The shoot
system is composed of stem, leaves, and
flowers. The stems hold and orient the
leaves to the sun, which allow the leaves
to conduct photosynthesis. The flowers
Root and shoot systems in a
are shoots that have been modified for eudicot
reproduction. Shoots are composed of
phytomers, which are functional units that consist of a node carrying
one or more leaves, internode, and one or more buds.
:
A plant body has two basic patterns (apical–basal and radial axes)
that been established during embryogenesis.[201] Cells and tissues
are arranged along the apical-basal axis from root to shoot whereas
the three tissue systems (dermal, ground, and vascular) that make
up a plant's body are arranged concentrically around its radial axis.
[201] The dermal tissue system forms the epidermis (or outer

covering) of a plant, which is usually a single cell layer that consists


of cells that have differentiated into three specialized structures:
stomata for gas exchange in leaves, trichomes (or leaf hair) for
protection against insects and solar radiation, and root hairs for
increased surface areas and absorption of water and nutrients. The
ground tissue makes up virtually all the tissue that lies between the
dermal and vascular tissues in the shoots and roots. It consists of
three cell types: Parenchyma, collenchyma, and sclerenchyma cells.
Finally, the vascular tissues are made up of two constituent tissues:
xylem and phloem. The xylem is made up of two conducting cells
called tracheids and vessel elements whereas the phloem is
characterized by the presence of sieve tube elements and
companion cells.[201]

Plant nutrition and transport

Further information: Vascular plant § Nutrient distribution

Like all other organisms, plants are


primarily made up of water and other
molecules containing elements that are
essential to life.[202] The absence of
specific nutrients (or essential elements),
many of which have been identified in
hydroponic experiments, can disrupt plant
growth and reproduction. The majority of
The xylem (blue) transports
plants are able to obtain these nutrients water and minerals from the
:
from solutions that surrounds their roots in roots upwards whereas the
phloem (orange) transports
the soil.[202] Continuous leaching and carbohydrates between organs.
harvesting of crops can deplete the soil of
its nutrients, which can be restored with the use of fertilizers.
Carnivorous plants such as Venus flytraps are able to obtain
nutrients by digesting other arthropods whereas parasitic plants
such as mistletoes can parasitize other plants for water and
nutrients.

Plants need water to conduct photosynthesis, transport solutes


between organs, cool their leaves by evaporation, and maintain
internal pressures that support their bodies.[202] Water is able to
diffuse in and out of plant cells by osmosis. The direction of water
movement across a semipermeable membrane is determined by the
water potential across that membrane.[202] Water is able to diffuse
across a root cell's membrane through aquaporins whereas solutes
are transported across by the membrane by ion channels and
pumps. In vascular plants, water and solutes are able to enter the
xylem, a vascular tissue, by way of an apoplast and symplast. Once
in the xylem, the water and minerals are distributed upward by
transpiration from the soil to the aerial parts of the plant.[175][202] In
contrast, the phloem, another vascular tissue, distributes
carbohydrates (e.g., sucrose) and other solutes such as hormones
by translocation from a source (e.g., mature leaf or root) in which
they were produced to a sink (e.g., root, flower, or developing fruit) in
which they will be used and stored.[202] Sources and sinks can
switch roles, depending on the amount of carbohydrates
accumulated or mobilized for the nourishment of other organs.

Plant development

Plant development is regulated by environmental cues and the


plant's own receptors, hormones, and genome.[203] Morever, they
:
have several characteristics that allow them to obtain resources for
growth and reproduction such as meristems, post-embryonic organ
formation, and differential growth.

Development begins with a seed, which is an embryonic plant


enclosed in a protective outer covering. Most plant seeds are usually
dormant, a condition in which the seed's normal activity is
suspended.[203] Seed dormancy may last may last weeks, months,
years, and even centuries. Dormancy is broken once conditions are
favorable for growth, and the seed will begin to sprout, a process
called germination. Imbibition is the first step in germination,
whereby water is absorbed by the seed. Once water is absorbed, the
seed undergoes metabolic changes whereby enzymes are activated
and RNA and proteins are synthesized. Once the seed germinates, it
obtains carbohydrates, amino acids, and small lipids that serve as
building blocks for its development. These monomers are obtained
from the hydrolysis of starch, proteins, and lipids that are stored in
either the cotyledons or endosperm. Germination is completed once
embryonic roots called radicle have emerged from the seed coat. At
this point, the developing plant is called a seedling and its growth is
regulated by its own photoreceptor proteins and hormones.[203]

Unlike animals in which growth is determinate, i.e., ceases when the


adult state is reached, plant growth is indeterminate as it is an open-
ended process that could potentially be lifelong.[201] Plants grow in
two ways: primary and secondary. In primary growth, the shoots and
roots are formed and lengthened. The apical meristem produces the
primary plant body, which can be found in all seed plants. During
secondary growth, the thickness of the plant increases as the lateral
meristem produces the secondary plant body, which can be found in
woody eudicots such as trees and shrubs. Monocots do not go
through secondary growth.[201] The plant body is generated by a
hierarchy of meristems. The apical meristems in the root and shoot
:
systems give rise to primary meristems (protoderm, ground
meristem, and procambium), which in turn, give rise to the three
tissue systems (dermal, ground, and vascular).

Plant reproduction

Further information: Plant reproduction

Most angiosperms (or flowering plants)


engage in sexual reproduction.[204] Their
flowers are organs that facilitate
reproduction, usually by providing a
mechanism for the union of sperm with Reproduction and development
eggs. Flowers may facilitate two types of in sporophytes
pollination: self-pollination and cross-pollination. Self-pollination
occurs when the pollen from the anther is deposited on the stigma of
the same flower, or another flower on the same plant. Cross-
pollination is the transfer of pollen from the anther of one flower to
the stigma of another flower on a different individual of the same
species. Self-pollination happened in flowers where the stamen and
carpel mature at the same time, and are positioned so that the pollen
can land on the flower's stigma. This pollination does not require an
investment from the plant to provide nectar and pollen as food for
pollinators.[205]

Plant responses

Further information: Plant perception (physiology) and Plant defense


against herbivory § Chemical defenses

Like animals, plants produce hormones in one part of its body to


signal cells in another part to respond. The ripening of fruit and loss
of leaves in the winter are controlled in part by the production of the
gas ethylene by the plant. Stress from water loss, changes in air
:
chemistry, or crowding by other plants can lead to changes in the
way a plant functions. These changes may be affected by genetic,
chemical, and physical factors.

To function and survive, plants produce a wide array of chemical


compounds not found in other organisms. Because they cannot
move, plants must also defend themselves chemically from
herbivores, pathogens and competition from other plants. They do
this by producing toxins and foul-tasting or smelling chemicals.
Other compounds defend plants against disease, permit survival
during drought, and prepare plants for dormancy, while other
compounds are used to attract pollinators or herbivores to spread
ripe seeds.

Many plant organs contain different types of photoreceptor proteins,


each of which reacts very specifically to certain wavelengths of light.
[206] The photoreceptor proteins relay information such as whether it

is day or night, duration of the day, intensity of light available, and the
source of light. Shoots generally grow towards light, while roots grow
away from it, responses known as phototropism and skototropism,
respectively. They are brought about by light-sensitive pigments like
phototropins and phytochromes and the plant hormone auxin.[207]
Many flowering plants bloom at the appropriate time because of
light-sensitive compounds that respond to the length of the night, a
phenomenon known as photoperiodism.

In addition to light, plants can respond to other types of stimuli. For


instance, plants can sense the direction of gravity to orient
themselves correctly. They can respond to mechanical stimulation.
[208]

Animal form and function


:
General features

Further information: Anatomy and Physiology

The cells in each animal body are bathed


in interstitial fluid, which make up the cell's
environment. This fluid and all its
characteristics (e.g., temperature, ionic
composition) can be described as the Negative feedback is necessary
animal's internal environment, which is in for maintaining homeostasis such
as keeping body temperature
contrast to the external environment that
constant.
encompasses the animal's outside world.
[209] Animals can be classified as either regulators or conformers.

Animals such as mammals and birds are regulators as they are able
to maintain a constant internal environment such as body
temperature despite their environments changing. These animals are
also described as homeotherms as they exhibit thermoregulation by
keeping their internal body temperature constant. In contrast,
animals such as fishes and frogs are conformers as they adapt their
internal environment (e.g., body temperature) to match their external
environments. These animals are also described as poikilotherms or
ectotherms as they allow their body temperatures to match their
external environments. In terms of energy, regulation is more costly
than conformity as an animal expands more energy to maintain a
constant internal environment such as increasing its basal metabolic
rate, which is the rate of energy consumption.[209] Similarly,
homeothermy is more costly than poikilothermy. Homeostasis is the
stability of an animal's internal environment, which is maintained by
negative feedback loops.[209][210]

The body size of terrestrial animals vary across different species but
their use of energy does not scale linearly according to their size.
[209] Mice, for example, are able to consume three times more food
:
than rabbits in proportion to their weights as the basal metabolic rate
per unit weight in mice is greater than in rabbits.[209] Physical activity
can also increase an animal's metabolic rate. When an animal runs,
its metabolic rate increases linearly with speed.[209] However, the
relationship is non-linear in animals that swim or fly. When a fish
swims faster, it encounters greater water resistance and so its
metabolic rates increases exponential.[209] Alternatively, the
relationship of flight speeds and metabolic rates is U-shaped in
birds.[209] At low flight speeds, a bird must maintain a high metabolic
rates to remain airborne. As it speeds up its flight, its metabolic rate
decreases with the aid of air rapidly flows over its wings. However, as
it increases in its speed even further, its high metabolic rates rises
again due to the increased effort associated with rapid flight speeds.
Basal metabolic rates can be measured based on an animal's rate of
heat production.

Water and salt balance

Further information: Osmoregulation and Urinary system

An animal's body fluids have three


properties: osmotic pressure, ionic
composition, and volume.[211] Osmotic
pressures determine the direction of the
Diffusion of water and ions in and
diffusion of water (or osmosis), which out of a freshwater fish
moves from a region where osmotic
pressure (total solute concentration) is low to a region where
osmotic pressure (total solute concentration) is high. Aquatic animals
are diverse with respect to their body fluid compositions and their
environments. For example, most invertebrate animals in the ocean
have body fluids that are isosmotic with seawater. In contrast, ocean
bony fishes have body fluids that are hyposmotic to seawater. Finally,
freshwater animals have body fluids that are hyperosmotic to fresh
:
water. Typical ions that can be found in an animal's body fluids are
sodium, potassium, calcium, and chloride. The volume of body fluids
can be regulated by excretion. Vertebrate animals have kidneys,
which are excretory organs made up of tiny tubular structures called
nephrons, which make urine from blood plasma. The kidneys'
primary function is to regulate the composition and volume of blood
plasma by selectively removing material from the blood plasma itself.
The ability of xeric animals such as kangaroo rats to minimize water
loss by producing urine that is 10–20 times concentrated than their
blood plasma allows them to adapt in desert environments that
receive very little precipitation.[211]

Nutrition and digestion

Further information: Nutrition

Animals are heterotrophs as they feed on


other organisms to obtain energy and
organic compounds.[212] They are able to
obtain food in three major ways such as
targeting visible food objects, collecting
tiny food particles, or depending on
microbes for critical food needs. The
amount of energy stored in food can be
quantified based on the amount of heat
(measured in calories or kilojoules)
Different digestive systems in
emitted when the food is burnt in the marine fishes
presence of oxygen. If an animal were to
consume food that contains an excess amount of chemical energy, it
will store most of that energy in the form of lipids for future use and
some of that energy as glycogen for more immediate use (e.g.,
meeting the brain's energy needs).[212] The molecules in food are
chemical building blocks that are needed for growth and
:
development. These molecules include nutrients such as
carbohydrates, fats, and proteins. Vitamins and minerals (e.g.,
calcium, magnesium, sodium, and phosphorus) are also essential.
The digestive system, which typically consist of a tubular tract that
extends from the mouth to the anus, is involved in the breakdown (or
digestion) of food into small molecules as it travels down
peristaltically through the gut lumen shortly after it has been
ingested. These small food molecules are then absorbed into the
blood from the lumen, where they are then distributed to the rest of
the body as building blocks (e.g., amino acids) or sources of energy
(e.g., glucose).[212]

In addition to their digestive tracts, vertebrate animals have


accessory glands such as a liver and pancreas as part of their
digestive systems.[212] The processing of food in these animals
begins in the foregut, which includes the mouth, esophagus, and
stomach. Mechanical digestion of food starts in the mouth with the
esophagus serving as a passageway for food to reach the stomach,
where it is stored and disintegrated (by the stomach's acid) for
further processing. Upon leaving the stomach, food enters into the
midgut, which is the first part of the intestine (or small intestine in
mammals) and is the principal site of digestion and absorption. Food
that does not get absorbed are stored as indigestible waste (or
feces) in the hindgut, which is the second part of the intestine (or
large intestine in mammals). The hindgut then completes the
reabsorption of needed water and salt prior to eliminating the feces
from the rectum.[212]

Breathing

The respiratory system consists of specific organs and structures


used for gas exchange in animals. The anatomy and physiology that
make this happen varies greatly, depending on the size of the
:
organism, the environment in which it lives
and its evolutionary history. In land animals
the respiratory surface is internalized as
linings of the lungs.[213] Gas exchange in
the lungs occurs in millions of small air
sacs; in mammals and reptiles these are
called alveoli, and in birds they are known
Respiratory system in a bird
as atria. These microscopic air sacs have a
very rich blood supply, thus bringing the air into close contact with
the blood.[214] These air sacs communicate with the external
environment via a system of airways, or hollow tubes, of which the
largest is the trachea, which branches in the middle of the chest into
the two main bronchi. These enter the lungs where they branch into
progressively narrower secondary and tertiary bronchi that branch
into numerous smaller tubes, the bronchioles. In birds the
bronchioles are termed parabronchi. It is the bronchioles, or
parabronchi that generally open into the microscopic alveoli in
mammals and atria in birds. Air has to be pumped from the
environment into the alveoli or atria by the process of breathing,
which involves the muscles of respiration.

Circulation

A circulatory system usually consists of a


muscular pump such as a heart, a fluid
(blood), and system of blood vessels that
deliver it.[215][216] Its principal function is to
transport blood and other substances to
and from cells and tissues. There are two Circulatory systems in
types of circulatory systems: open and arthropods, fish, reptiles, and
closed. In open circulatory systems, blood birds/mammals
exits blood vessels as it circulates throughout the body whereas in
closed circulatory system, blood is contained within the blood
:
vessels as it circulates. Open circulatory systems can be observed in
invertebrate animals such as arthropods (e.g., insects, spiders, and
lobsters) whereas closed circulatory systems can be found in
vertebrate animals such as fishes, amphibians, and mammals.
Circulation in animals occur between two types of tissues: systemic
tissues and breathing (or pulmonary) organs.[215] Systemic tissues
are all the tissues and organs that make up an animal's body other
than its breathing organs. Systemic tissues take up oxygen but adds
carbon dioxide to the blood whereas a breathing organs takes up
carbon dioxide but add oxygen to the blood.[217] In birds and
mammals, the systemic and pulmonary systems are connected in
series.

In the circulatory system, blood is important because it is the means


by which oxygen, carbon dioxide, nutrients, hormones, agents of
immune system, heat, wastes, and other commodities are
transported.[215] In annelids such as earthworms and leeches, blood
is propelled by peristaltic waves of contractions of the heart muscles
that make up the blood vessels. Other animals such as crustaceans
(e.g., crayfish and lobsters), have more than one heart to propel
blood throughout their bodies. Vertebrate hearts are multichambered
and are able to pump blood when their ventricles contract at each
cardiac cycle, which propels blood through the blood vessels.[215]
Although vertebrate hearts are myogenic, their rate of contraction
(or heart rate) can be modulated by neural input from the body's
autonomic nervous system.

Muscle and movement

Further information: Muscle contraction

In vertebrates, the muscular system consists of skeletal, smooth and


cardiac muscles. It permits movement of the body, maintains posture
:
and circulates blood throughout the body.
[218] Together with the skeletal system, it

forms the musculoskeletal system, which


is responsible for the movement of
vertebrate animals.[219] Skeletal muscle
contractions are neurogenic as they
require synaptic input from motor neurons.
A single motor neuron is able to innervate Asynchronous muscles power
flight in most insects. a: Wings b:
multiple muscle fibers, thereby causing Wing joint c: Dorsoventral
the fibers to contract at the same time. muscles power upstrokes d:
Dorsolongitudinal muscles power
Once innervated, the protein filaments
downstrokes.
within each skeletal muscle fiber slide past
each other to produce a contraction, which is explained by the
sliding filament theory. The contraction produced can be described
as a twitch, summation, or tetanus, depending on the frequency of
action potentials. Unlike skeletal muscles, contractions of smooth
and cardiac muscles are myogenic as they are initiated by the
smooth or heart muscle cells themselves instead of a motor neuron.
Nevertheless, the strength of their contractions can be modulated by
input from the autonomic nervous system. The mechanisms of
contraction are similar in all three muscle tissues.

In invertebrates such as earthworms and leeches, circular and


longitudinal muscles cells form the body wall of these animals and
are responsible for their movement.[220] In an earthworm that is
moving through a soil, for example, contractions of circular and
longitudinal muscles occur reciprocally while the coelomic fluid
serves as a hydroskeleton by maintaining turgidity of the earthworm.
[221] Other animals such as mollusks, and nematodes, possess

obliquely striated muscles, which contain bands of thick and thin


filaments that are arranged helically rather than transversely, like in
vertebrate skeletal or cardiac muscles.[222] Advanced insects such
:
as wasps, flies, bees, and beetles possess asynchronous muscles
that constitute the flight muscles in these animals.[222] These flight
muscles are often called fibrillar muscles because they contain
myofibrils that are thick and conspicuous.[223]

Nervous system

Further information: Neuroscience and Neuroethology

Most multicellular animals have nervous


systems[225] that allow them to sense from
and respond to their environments. A
nervous system is a network of cells that
processes sensory information and
generates behaviors. At the cellular level,
the nervous system is defined by the Mouse pyramidal neurons
(green) and GABAergic neurons
presence of neurons, which are cells (red)[224]
specialized to handle information. [226]

They can transmit or receive information at sites of contacts called


synapses.[226] More specifically, neurons can conduct nerve
impulses (or action potentials) that travel along their thin fibers
called axons, which can then be transmitted directly to a neighboring
cell through electrical synapses or cause chemicals called
neurotransmitters to be released at chemical synapses. According to
the sodium theory, these action potentials can be generated by the
increased permeability of the neuron's cell membrane to sodium
ions.[227] Cells such as neurons or muscle cells may be excited or
inhibited upon receiving a signal from another neuron. The
connections between neurons can form neural pathways, neural
circuits, and larger networks that generate an organism's perception
of the world and determine its behavior. Along with neurons, the
nervous system contains other specialized cells called glia or glial
cells, which provide structural and metabolic support.
:
In vertebrates, the nervous system comprises the central nervous
system (CNS), which includes the brain and spinal cord, and the
peripheral nervous system (PNS), which consists of nerves that
connect the CNS to every other part of the body. Nerves that
transmit signals from the CNS are called motor nerves or efferent
nerves, while those nerves that transmit information from the body
to the CNS are called sensory nerves or afferent nerves. Spinal
nerves are mixed nerves that serve both functions. The PNS is
divided into three separate subsystems, the somatic, autonomic, and
enteric nervous systems. Somatic nerves mediate voluntary
movement. The autonomic nervous system is further subdivided into
the sympathetic and the parasympathetic nervous systems. The
sympathetic nervous system is activated in cases of emergencies to
mobilize energy, while the parasympathetic nervous system is
activated when organisms are in a relaxed state. The enteric nervous
system functions to control the gastrointestinal system. Both
autonomic and enteric nervous systems function involuntarily.
Nerves that exit directly from the brain are called cranial nerves while
those exiting from the spinal cord are called spinal nerves.

Many animals have sense organs that can detect their environment.
These sense organs contain sensory receptors, which are sensory
neurons that convert stimuli into electrical signals.[228]
Mechanoreceptors, for example, which can be found in skin, muscle,
and hearing organs, generate action potentials in response to
changes in pressures.[228][229] Photoreceptor cells such as rods and
cones, which are part of the vertebrate retina, can respond to
specific wavelengths of light.[228][229] Chemoreceptors detect
chemicals in the mouth (taste) or in the air (smell).[229]

Hormonal control

Further information: Endocrinology


:
Hormones are signaling molecules transported in the blood to
distant organs to regulate their function.[230][231] Hormones are
secreted by internal glands that are part of an animal's endocrine
system. In vertebrates, the hypothalamus is the neural control center
for all endocrine systems. In humans specifically, the major
endocrine glands are the thyroid gland and the adrenal glands. Many
other organs that are part of other body systems have secondary
endocrine functions, including bone, kidneys, liver, heart and
gonads. For example, kidneys secrete the endocrine hormone
erythropoietin. Hormones can be amino acid complexes, steroids,
eicosanoids, leukotrienes, or prostaglandins.[232] The endocrine
system can be contrasted to both exocrine glands, which secrete
hormones to the outside of the body, and paracrine signaling
between cells over a relatively short distance. Endocrine glands have
no ducts, are vascular, and commonly have intracellular vacuoles or
granules that store their hormones. In contrast, exocrine glands,
such as salivary glands, sweat glands, and glands within the
gastrointestinal tract, tend to be much less vascular and have ducts
or a hollow lumen.

Animal reproduction

Animals can reproduce in one of two ways:


asexual and sexual. Nearly all animals
engage in some form of sexual
reproduction.[233] They produce haploid
gametes by meiosis. The smaller, motile
gametes are spermatozoa and the larger, Sexual reproduction in
non-motile gametes are ova.[234] These dragonflies
fuse to form zygotes,[235] which develop
via mitosis into a hollow sphere, called a blastula. In sponges,
blastula larvae swim to a new location, attach to the seabed, and
develop into a new sponge.[236] In most other groups, the blastula
:
undergoes more complicated rearrangement.[237] It first invaginates
to form a gastrula with a digestive chamber and two separate germ
layers, an external ectoderm and an internal endoderm.[238] In most
cases, a third germ layer, the mesoderm, also develops between
them.[239] These germ layers then differentiate to form tissues and
organs.[240] Some animals are capable of asexual reproduction,
which often results in a genetic clone of the parent. This may take
place through fragmentation; budding, such as in Hydra and other
cnidarians; or parthenogenesis, where fertile eggs are produced
without mating, such as in aphids.[241][242]
Animal development

Further information: Developmental biology and Embryology

Animal development begins with the


formation of a zygote that results from the
fusion of a sperm and egg during
fertilization.[243] The zygote undergoes a
rapid multiple rounds of mitotic cell period
of cell divisions called cleavage, which Cleavage in zebrafish embryo
forms a ball of similar cells called a
blastula. Gastrulation occurs, whereby morphogenetic movements
convert the cell mass into a three germ layers that comprise the
ectoderm, mesoderm and endoderm.

The end of gastrulation signals the beginning of organogenesis,


whereby the three germ layers form the internal organs of the
organism.[244] The cells of each of the three germ layers undergo
differentiation, a process where less-specialized cells become more-
specialized through the expression of a specific set of genes.
Cellular differentiation is influenced by extracellular signals such as
growth factors that are exchanged to adjacent cells, which is called
juxtracrine signaling, or to neighboring cells over short distances,
:
which is called paracrine signaling.[245][246] Intracellular signals
consist of a cell signaling itself (autocrine signaling), also play a role
in organ formation. These signaling pathways allows for cell
rearrangement and ensures that organs form at specific sites within
the organism.[244][247]

Immune system

Further information: Immunology

The immune system is a network of


biological processes that detects and
responds to a wide variety of pathogens.
Many species have two major subsystems
of the immune system. The innate immune
system provides a preconfigured response
Processes in the primary immune
to broad groups of situations and stimuli. response
The adaptive immune system provides a
tailored response to each stimulus by learning to recognize
molecules it has previously encountered. Both use molecules and
cells to perform their functions.

Nearly all organisms have some kind of immune system. Bacteria


have a rudimentary immune system in the form of enzymes that
protect against virus infections. Other basic immune mechanisms
evolved in ancient plants and animals and remain in their modern
descendants. These mechanisms include phagocytosis,
antimicrobial peptides called defensins, and the complement system.
Jawed vertebrates, including humans, have even more sophisticated
defense mechanisms, including the ability to adapt to recognize
pathogens more efficiently. Adaptive (or acquired) immunity creates
an immunological memory leading to an enhanced response to
subsequent encounters with that same pathogen. This process of
:
acquired immunity is the basis of vaccination.

Animal behavior

Further information: Ethology

Behaviors play a central a role in animals'


interaction with each other and with their
environment.[248] They are able to use their
muscles to approach one another, vocalize, seek
shelter, and migrate. An animal's nervous system
activates and coordinates its behaviors. Fixed
action patterns, for instance, are genetically
Brood parasites, such determined and stereotyped behaviors that occur
as the cuckoo, provide without learning.[248][249] These behaviors are
a supernormal
stimulus to the
under the control of the nervous system and can
parenting species. be quite elaborate.[248] Examples include the
pecking of kelp gull chicks at the red dot on their
mother's beak. Other behaviors that have emerged as a result of
natural selection include foraging, mating, and altruism.[250] In
addition to evolved behavior, animals have evolved the ability to learn
by modifying their behaviors as a result of early individual
experiences.[248]

Ecology

Ecosystems

Ecology is the study of the distribution and abundance of life, the


interaction between organisms and their environment.[251] The
community of living (biotic) organisms in conjunction with the
nonliving (abiotic) components (e.g., water, light, radiation,
temperature, humidity, atmosphere, acidity, and soil) of their
:
environment is called an ecosystem.[252]
[253][254] These biotic and abiotic

components are linked together through


nutrient cycles and energy flows.[255]
Energy from the sun enters the system
through photosynthesis and is
incorporated into plant tissue. By feeding
on plants and on one another, animals play
an important role in the movement of Terrestrial biomes are shaped by
temperature and precipitation.
matter and energy through the system.
They also influence the quantity of plant and microbial biomass
present. By breaking down dead organic matter, decomposers
release carbon back to the atmosphere and facilitate nutrient cycling
by converting nutrients stored in dead biomass back to a form that
can be readily used by plants and other microbes.[256]

The Earth's physical environment is shaped by solar energy and


topography.[254] The amount of solar energy input varies in space
and time due to the spherical shape of the Earth and its axial tilt.
Variation in solar energy input drives weather and climate patterns.
Weather is the day-to-day temperature and precipitation activity,
whereas climate is the long-term average of weather, typically
averaged over a period of 30 years.[257][258] Variation in topography
also produces environmental heterogeneity. On the windward side of
a mountain, for example, air rises and cools, with water changing
from gaseous to liquid or solid form, resulting in precipitation such as
rain or snow.[254] As a result, wet environments allow for lush
vegetation to grow. In contrast, conditions tend to be dry on the
leeward side of a mountain due to the lack of precipitation as air
descends and warms, and moisture remains as water vapor in the
atmosphere. Temperature and precipitation are the main factors that
shape terrestrial biomes.
:
Populations

Further information: Population ecology

A population is the number of organisms


of the same species that occupy an area
and reproduce from generation to
generation.[259][260][261][262][263] Its
abundance can be measured using
population density, which is the number of
Reaching carrying capacity
individuals per unit area (e.g., land or tree) through a logistic growth curve
or volume (e.g., sea or air).[259] Given that
it is usually impractical to count every individual within a large
population to determine its size, population size can be estimated by
multiplying population density by the area or volume. Population
growth during short-term intervals can be determined using the
population growth rate equation, which takes into consideration
birth, death, and immigration rates. In the longer term, the
exponential growth of a population tends to slow down as it reaches
its carrying capacity, which can be modeled using the logistic
equation.[260] The carrying capacity of an environment is the
maximum population size of a species that can be sustained by that
specific environment, given the food, habitat, water, and other
resources that are available.[264] The carrying capacity of a
population can be affected by changing environmental conditions
such as changes in the availability resources and the cost of
maintaining them. In human populations, new technologies such as
the Green revolution have helped increase the Earth's carrying
capacity for humans over time, which has stymied the attempted
predictions of impending population decline, the famous of which
was by Thomas Malthus in the 18th century.[259]

Communities
:
A (a) trophic pyramid and a (b) simplified food web. The
trophic pyramid represents the biomass at each level.
[265]

A community is a group of populations of two or more different


species occupying the same geographical area at the same time. A
biological interaction is the effect that a pair of organisms living
together in a community have on each other. They can be either of
the same species (intraspecific interactions), or of different species
(interspecific interactions). These effects may be short-term, like
pollination and predation, or long-term; both often strongly influence
the evolution of the species involved. A long-term interaction is
called a symbiosis. Symbioses range from mutualism, beneficial to
both partners, to competition, harmful to both partners.[266]

Every species participates as a consumer, resource, or both in


consumer–resource interactions, which form the core of food chains
or food webs.[267] There are different trophic levels within any food
web, with the lowest level being the primary producers (or
autotrophs) such as plants and algae that convert energy and
inorganic material into organic compounds, which can then be used
by the rest of the community.[62][268][269] At the next level are the
heterotrophs, which are the species that obtain energy by breaking
apart organic compounds from other organisms.[267] Heterotrophs
that consume plants are primary consumers (or herbivores) whereas
heterotrophs that consume herbivores are secondary consumers (or
carnivores). And those that eat secondary consumers are tertiary
consumers and so on. Omnivorous heterotrophs are able to
:
consume at multiple levels. Finally, there are decomposers that feed
on the waste products or dead bodies of organisms.[267]

On average, the total amount of energy incorporated into the


biomass of a trophic level per unit of time is about one-tenth of the
energy of the trophic level that it consumes. Waste and dead
material used by decomposers as well as heat lost from metabolism
make up the other ninety percent of energy that is not consumed by
the next trophic level.[270]

Biosphere

In the global ecosystem (or biosphere),


matter exist as different interacting
compartments, which can be biotic or
abiotic as well as accessible or
inaccessible, depending on their forms
and locations.[272] For example, matter
from terrestrial autotrophs are both
biotic and accessible to other
organisms whereas the matter in rocks Fast carbon cycle showing the
movement of carbon between land,
and minerals are abiotic and atmosphere, and oceans in billions of
inaccessible. A biogeochemical cycle istons per year. Yellow numbers are
a pathway by which specific elements natural fluxes, red are human
contributions, white are stored
of matter are turned over or moved carbon. Effects of the slow carbon
through the biotic (biosphere) and the cycle, such as volcanic and tectonic
activity, are not included.[271]
abiotic (lithosphere, atmosphere, and
hydrosphere) compartments of Earth. There are biogeochemical
cycles for nitrogen, carbon, and water. In some cycles there are
reservoirs where a substance remains or is sequestered for a long
period of time.

Climate change includes both global warming driven by human-


:
induced emissions of greenhouse gases and the resulting large-
scale shifts in weather patterns. Though there have been previous
periods of climatic change, since the mid-20th century humans have
had an unprecedented impact on Earth's climate system and caused
change on a global scale.[273] The largest driver of warming is the
emission of greenhouse gases, of which more than 90% are carbon
dioxide and methane.[274] Fossil fuel burning (coal, oil, and natural
gas) for energy consumption is the main source of these emissions,
with additional contributions from agriculture, deforestation, and
manufacturing.[275] Temperature rise is accelerated or tempered by
climate feedbacks, such as loss of sunlight-reflecting snow and ice
cover, increased water vapor (a greenhouse gas itself), and changes
to land and ocean carbon sinks.

Conservation

Conservation biology is the study of the


conservation of Earth's biodiversity with
the aim of protecting species, their
habitats, and ecosystems from excessive
rates of extinction and the erosion of biotic
interactions.[276][277][278] It is concerned
Efforts are made to preserve the
with factors that influence the natural characteristics of
maintenance, loss, and restoration of Hopetoun Falls, Australia, without
biodiversity and the science of sustaining affecting visitors' access.
evolutionary processes that engender genetic, population, species,
and ecosystem diversity.[279][280][281][282] The concern stems from
estimates suggesting that up to 50% of all species on the planet will
disappear within the next 50 years,[283] which has contributed to
poverty, starvation, and will reset the course of evolution on this
planet.[284][285] Biodiversity affects the functioning of ecosystems,
which provide a variety of services upon which people depend.
:
Conservation biologists research and educate on the trends of
biodiversity loss, species extinctions, and the negative effect these
are having on our capabilities to sustain the well-being of human
society. Organizations and citizens are responding to the current
biodiversity crisis through conservation action plans that direct
research, monitoring, and education programs that engage concerns
at local through global scales.[286][279][280][281]

See also
Biology in fiction
Glossary of biology
List of biological websites
List of biologists
List of biology journals
List of biology topics
List of life sciences
List of omics topics in biology
National Association of Biology Teachers
Outline of biology
Periodic table of life sciences in Tinbergen's four questions
Reproduction
Science tourism
Terminology of biology

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doi:10.1126/science.1101101. PMID 15361627. S2CID 30713492.
284. ^ Millennium Ecosystem Assessment (2005). Ecosystems and
Human Well-being: Biodiversity Synthesis. World Resources
Institute, Washington, D.C.[1]
285. ^ Jackson, J. B. C. (2008). "Ecological extinction and evolution
in the brave new ocean". Proceedings of the National Academy
of Sciences. 105 (Suppl 1): 11458–65.
Bibcode:2008PNAS..10511458J. doi:10.1073/pnas.0802812105.
PMC 2556419. PMID 18695220.
286. ^ Soule, Michael E. (1986). Conservation Biology: The Science
of Scarcity and Diversity. Sinauer Associates. p. 584. ISBN 978-
0-87893-795-0.
:
Further reading
Further information: Bibliography of biology

Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P


(2002). Molecular Biology of the Cell (4th ed.). Garland.
ISBN 978-0-8153-3218-3. OCLC 145080076.
Begon M, Townsend CR, Harper JL (2005). Ecology: From
Individuals to Ecosystems (4th ed.). Blackwell Publishing
Limited. ISBN 978-1-4051-1117-1. OCLC 57639896.
Campbell N (2004). Biology (7th ed.). Benjamin-Cummings
Publishing Company. ISBN 978-0-8053-7146-8.
OCLC 71890442.
Colinvaux P (1979). Why Big Fierce Animals are Rare: An
Ecologist's Perspective (reissue ed.). Princeton University
Press. ISBN 978-0-691-02364-9. OCLC 10081738.
Mayr, Ernst (1982). The Growth of Biological Thought: Diversity,
Evolution, and Inheritance. Harvard University Press. ISBN 978-
0-674-36446-2. Archived from the original on 2015-10-03.
Retrieved 2015-06-27.
Hoagland M (2001). The Way Life Works (reprint ed.). Jones
and Bartlett Publishers inc. ISBN 978-0-7637-1688-2.
OCLC 223090105.
Janovy, John (2004). On Becoming a Biologist (2nd ed.). Bison
Books. ISBN 978-0-8032-7620-8. OCLC 55138571.
Johnson, George B. (2005). Biology, Visualizing Life. Holt,
Rinehart, and Winston. ISBN 978-0-03-016723-2.
OCLC 36306648.
Tobin, Allan; Dusheck, Jennie (2005). Asking About Life
(3rd ed.). Belmont, Calif.: Wadsworth. ISBN 978-0-534-40653-
0.
:
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