12.3.2: Components and Structure - Biology

Skills to Develop

  • Understand the fluid mosaic model of cell membranes
  • Describe the functions of phospholipids, proteins, and carbohydrates in membranes
  • Discuss membrane fluidity

A cell’s plasma membrane defines the cell, outlines its borders, and determines the nature of its interaction with its environment (see Table 5.1.1 for a summary). Cells exclude some substances, take in others, and excrete still others, all in controlled quantities. The plasma membrane must be very flexible to allow certain cells, such as red blood cells and white blood cells, to change shape as they pass through narrow capillaries. These are the more obvious functions of a plasma membrane. In addition, the surface of the plasma membrane carries markers that allow cells to recognize one another, which is vital for tissue and organ formation during early development, and which later plays a role in the “self” versus “non-self” distinction of the immune response.

Among the most sophisticated functions of the plasma membrane is the ability to transmit signals by means of complex, integral proteins known as receptors. These proteins act both as receivers of extracellular inputs and as activators of intracellular processes. These membrane receptors provide extracellular attachment sites for effectors like hormones and growth factors, and they activate intracellular response cascades when their effectors are bound. Occasionally, receptors are hijacked by viruses (HIV, human immunodeficiency virus, is one example) that use them to gain entry into cells, and at times, the genes encoding receptors become mutated, causing the process of signal transduction to malfunction with disastrous consequences.

Fluid Mosaic Model

The existence of the plasma membrane was identified in the 1890s, and its chemical components were identified in 1915. The principal components identified at that time were lipids and proteins. The first widely accepted model of the plasma membrane’s structure was proposed in 1935 by Hugh Davson and James Danielli; it was based on the “railroad track” appearance of the plasma membrane in early electron micrographs. They theorized that the structure of the plasma membrane resembles a sandwich, with protein being analogous to the bread, and lipids being analogous to the filling. In the 1950s, advances in microscopy, notably transmission electron microscopy (TEM), allowed researchers to see that the core of the plasma membrane consisted of a double, rather than a single, layer. A new model that better explains both the microscopic observations and the function of that plasma membrane was proposed by S.J. Singer and Garth L. Nicolson in 1972.

The explanation proposed by Singer and Nicolson is called the fluid mosaic model. The model has evolved somewhat over time, but it still best accounts for the structure and functions of the plasma membrane as we now understand them. The fluid mosaic model describes the structure of the plasma membrane as a mosaic of components—including phospholipids, cholesterol, proteins, and carbohydrates—that gives the membrane a fluid character. Plasma membranes range from 5 to 10 nm in thickness. For comparison, human red blood cells, visible via light microscopy, are approximately 8 µm wide, or approximately 1,000 times wider than a plasma membrane. The membrane does look a bit like a sandwich (Figure (PageIndex{1})).

The principal components of a plasma membrane are lipids (phospholipids and cholesterol), proteins, and carbohydrates attached to some of the lipids and some of the proteins. A phospholipid is a molecule consisting of glycerol, two fatty acids, and a phosphate-linked head group. Cholesterol, another lipid composed of four fused carbon rings, is found alongside the phospholipids in the core of the membrane. The proportions of proteins, lipids, and carbohydrates in the plasma membrane vary with cell type, but for a typical human cell, protein accounts for about 50 percent of the composition by mass, lipids (of all types) account for about 40 percent of the composition by mass, with the remaining 10 percent of the composition by mass being carbohydrates. However, the concentration of proteins and lipids varies with different cell membranes. For example, myelin, an outgrowth of the membrane of specialized cells that insulates the axons of the peripheral nerves, contains only 18 percent protein and 76 percent lipid. The mitochondrial inner membrane contains 76 percent protein and only 24 percent lipid. The plasma membrane of human red blood cells is 30 percent lipid. Carbohydrates are present only on the exterior surface of the plasma membrane and are attached to proteins, forming glycoproteins, or attached to lipids, forming glycolipids.


The main fabric of the membrane is composed of amphiphilic, phospholipid molecules. The hydrophilic or “water-loving” areas of these molecules (which look like a collection of balls in an artist’s rendition of the model) (Figure (PageIndex{1})) are in contact with the aqueous fluid both inside and outside the cell. Hydrophobic, or water-hating molecules, tend to be non-polar. They interact with other non-polar molecules in chemical reactions, but generally do not interact with polar molecules. When placed in water, hydrophobic molecules tend to form a ball or cluster. The hydrophilic regions of the phospholipids tend to form hydrogen bonds with water and other polar molecules on both the exterior and interior of the cell. Thus, the membrane surfaces that face the interior and exterior of the cell are hydrophilic. In contrast, the interior of the cell membrane is hydrophobic and will not interact with water. Therefore, phospholipids form an excellent two-layer cell membrane that separates fluid within the cell from the fluid outside of the cell.

A phospholipid molecule (Figure (PageIndex{2})) consists of a three-carbon glycerol backbone with two fatty acid molecules attached to carbons 1 and 2, and a phosphate-containing group attached to the third carbon. This arrangement gives the overall molecule an area described as its head (the phosphate-containing group), which has a polar character or negative charge, and an area called the tail (the fatty acids), which has no charge. The head can form hydrogen bonds, but the tail cannot. A molecule with this arrangement of a positively or negatively charged area and an uncharged, or non-polar, area is referred to as amphiphilic or “dual-loving.”

This characteristic is vital to the structure of a plasma membrane because, in water, phospholipids tend to become arranged with their hydrophobic tails facing each other and their hydrophilic heads facing out. In this way, they form a lipid bilayer—a barrier composed of a double layer of phospholipids that separates the water and other materials on one side of the barrier from the water and other materials on the other side. In fact, phospholipids heated in an aqueous solution tend to spontaneously form small spheres or droplets (called micelles or liposomes), with their hydrophilic heads forming the exterior and their hydrophobic tails on the inside (Figure (PageIndex{3})).


Proteins make up the second major component of plasma membranes. Integral proteins (some specialized types are called integrins) are, as their name suggests, integrated completely into the membrane structure, and their hydrophobic membrane-spanning regions interact with the hydrophobic region of the the phospholipid bilayer (Figure (PageIndex{1})). Single-pass integral membrane proteins usually have a hydrophobic transmembrane segment that consists of 20–25 amino acids. Some span only part of the membrane—associating with a single layer—while others stretch from one side of the membrane to the other, and are exposed on either side. Some complex proteins are composed of up to 12 segments of a single protein, which are extensively folded and embedded in the membrane (Figure (PageIndex{4})). This type of protein has a hydrophilic region or regions, and one or several mildly hydrophobic regions. This arrangement of regions of the protein tends to orient the protein alongside the phospholipids, with the hydrophobic region of the protein adjacent to the tails of the phospholipids and the hydrophilic region or regions of the protein protruding from the membrane and in contact with the cytosol or extracellular fluid.

Peripheral proteins are found on the exterior and interior surfaces of membranes, attached either to integral proteins or to phospholipids. Peripheral proteins, along with integral proteins, may serve as enzymes, as structural attachments for the fibers of the cytoskeleton, or as part of the cell’s recognition sites. These are sometimes referred to as “cell-specific” proteins. The body recognizes its own proteins and attacks foreign proteins associated with invasive pathogens.


Carbohydrates are the third major component of plasma membranes. They are always found on the exterior surface of cells and are bound either to proteins (forming glycoproteins) or to lipids (forming glycolipids) (Figure (PageIndex{1})). These carbohydrate chains may consist of 2–60 monosaccharide units and can be either straight or branched. Along with peripheral proteins, carbohydrates form specialized sites on the cell surface that allow cells to recognize each other. These sites have unique patterns that allow the cell to be recognized, much the way that the facial features unique to each person allow him or her to be recognized. This recognition function is very important to cells, as it allows the immune system to differentiate between body cells (called “self”) and foreign cells or tissues (called “non-self”). Similar types of glycoproteins and glycolipids are found on the surfaces of viruses and may change frequently, preventing immune cells from recognizing and attacking them.

These carbohydrates on the exterior surface of the cell—the carbohydrate components of both glycoproteins and glycolipids—are collectively referred to as the glycocalyx (meaning “sugar coating”). The glycocalyx is highly hydrophilic and attracts large amounts of water to the surface of the cell. This aids in the interaction of the cell with its watery environment and in the cell’s ability to obtain substances dissolved in the water. As discussed above, the glycocalyx is also important for cell identification, self/non-self determination, and embryonic development, and is used in cell-cell attachments to form tissues.

Evolution Connection

How Viruses Infect Specific OrgansGlycoprotein and glycolipid patterns on the surfaces of cells give many viruses an opportunity for infection. HIV and hepatitis viruses infect only specific organs or cells in the human body. HIV is able to penetrate the plasma membranes of a subtype of lymphocytes called T-helper cells, as well as some monocytes and central nervous system cells. The hepatitis virus attacks liver cells.

These viruses are able to invade these cells, because the cells have binding sites on their surfaces that are specific to and compatible with certain viruses (Figure (PageIndex{5})). Other recognition sites on the virus’s surface interact with the human immune system, prompting the body to produce antibodies. Antibodies are made in response to the antigens or proteins associated with invasive pathogens, or in response to foreign cells, such as might occur with an organ transplant. These same sites serve as places for antibodies to attach and either destroy or inhibit the activity of the virus. Unfortunately, these recognition sites on HIV change at a rapid rate because of mutations, making the production of an effective vaccine against the virus very difficult, as the virus evolves and adapts. A person infected with HIV will quickly develop different populations, or variants, of the virus that are distinguished by differences in these recognition sites. This rapid change of surface markers decreases the effectiveness of the person’s immune system in attacking the virus, because the antibodies will not recognize the new variations of the surface patterns. In the case of HIV, the problem is compounded by the fact that the virus specifically infects and destroys cells involved in the immune response, further incapacitating the host.

The mosaic characteristic of the membrane, described in the fluid mosaic model, helps to illustrate its nature. The integral proteins and lipids exist in the membrane as separate but loosely attached molecules. These resemble the separate, multicolored tiles of a mosaic picture, and they float, moving somewhat with respect to one another. The membrane is not like a balloon, however, that can expand and contract; rather, it is fairly rigid and can burst if penetrated or if a cell takes in too much water. However, because of its mosaic nature, a very fine needle can easily penetrate a plasma membrane without causing it to burst, and the membrane will flow and self-seal when the needle is extracted.

The mosaic characteristics of the membrane explain some but not all of its fluidity. There are two other factors that help maintain this fluid characteristic. One factor is the nature of the phospholipids themselves. In their saturated form, the fatty acids in phospholipid tails are saturated with bound hydrogen atoms. There are no double bonds between adjacent carbon atoms. This results in tails that are relatively straight. In contrast, unsaturated fatty acids do not contain a maximal number of hydrogen atoms, but they do contain some double bonds between adjacent carbon atoms; a double bond results in a bend in the string of carbons of approximately 30 degrees (Figure (PageIndex{2})).

Thus, if saturated fatty acids, with their straight tails, are compressed by decreasing temperatures, they press in on each other, making a dense and fairly rigid membrane. If unsaturated fatty acids are compressed, the “kinks” in their tails elbow adjacent phospholipid molecules away, maintaining some space between the phospholipid molecules. This “elbow room” helps to maintain fluidity in the membrane at temperatures at which membranes with saturated fatty acid tails in their phospholipids would “freeze” or solidify. The relative fluidity of the membrane is particularly important in a cold environment. A cold environment tends to compress membranes composed largely of saturated fatty acids, making them less fluid and more susceptible to rupturing. Many organisms (fish are one example) are capable of adapting to cold environments by changing the proportion of unsaturated fatty acids in their membranes in response to the lowering of the temperature.

Link to Learning

Visit this site to see animations of the fluidity and mosaic quality of membranes.

Animals have an additional membrane constituent that assists in maintaining fluidity. Cholesterol, which lies alongside the phospholipids in the membrane, tends to dampen the effects of temperature on the membrane. Thus, this lipid functions as a buffer, preventing lower temperatures from inhibiting fluidity and preventing increased temperatures from increasing fluidity too much. Thus, cholesterol extends, in both directions, the range of temperature in which the membrane is appropriately fluid and consequently functional. Cholesterol also serves other functions, such as organizing clusters of transmembrane proteins into lipid rafts.

Table (PageIndex{1}): The Components and Functions of the Plasma Membrane.
PhospholipidMain fabric of the membrane
CholesterolAttached between phospholipids and between the two phospholipid layers
Integral proteins (for example, integrins)Embedded within the phospholipid layer(s). May or may not penetrate through both layers
Peripheral proteinsOn the inner or outer surface of the phospholipid bilayer; not embedded within the phospholipids
Carbohydrates (components of glycoproteins and glycolipids)Generally attached to proteins on the outside membrane layer

Career Connection: Immunologist

The variations in peripheral proteins and carbohydrates that affect a cell’s recognition sites are of prime interest in immunology. These changes are taken into consideration in vaccine development. Many infectious diseases, such as smallpox, polio, diphtheria, and tetanus, were conquered by the use of vaccines.

Immunologists are the physicians and scientists who research and develop vaccines, as well as treat and study allergies or other immune problems. Some immunologists study and treat autoimmune problems (diseases in which a person’s immune system attacks his or her own cells or tissues, such as lupus) and immunodeficiencies, whether acquired (such as acquired immunodeficiency syndrome, or AIDS) or hereditary (such as severe combined immunodeficiency, or SCID). Immunologists are called in to help treat organ transplantation patients, who must have their immune systems suppressed so that their bodies will not reject a transplanted organ. Some immunologists work to understand natural immunity and the effects of a person’s environment on it. Others work on questions about how the immune system affects diseases such as cancer. In the past, the importance of having a healthy immune system in preventing cancer was not at all understood.

To work as an immunologist, a PhD or MD is required. In addition, immunologists undertake at least 2–3 years of training in an accredited program and must pass an examination given by the American Board of Allergy and Immunology. Immunologists must possess knowledge of the functions of the human body as they relate to issues beyond immunization, and knowledge of pharmacology and medical technology, such as medications, therapies, test materials, and surgical procedures.

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The modern understanding of the plasma membrane is referred to as the fluid mosaic model. The plasma membrane is composed of a bilayer of phospholipids, with their hydrophobic, fatty acid tails in contact with each other. The landscape of the membrane is studded with proteins, some of which span the membrane. Some of these proteins serve to transport materials into or out of the cell. Carbohydrates are attached to some of the proteins and lipids on the outward-facing surface of the membrane, forming complexes that function to identify the cell to other cells. The fluid nature of the membrane is due to temperature, the configuration of the fatty acid tails (some kinked by double bonds), the presence of cholesterol embedded in the membrane, and the mosaic nature of the proteins and protein-carbohydrate combinations, which are not firmly fixed in place. Plasma membranes enclose and define the borders of cells, but rather than being a static bag, they are dynamic and constantly in flux.

molecule possessing a polar or charged area and a nonpolar or uncharged area capable of interacting with both hydrophilic and hydrophobic environments
fluid mosaic model
describes the structure of the plasma membrane as a mosaic of components including phospholipids, cholesterol, proteins, glycoproteins, and glycolipids (sugar chains attached to proteins or lipids, respectively), resulting in a fluid character (fluidity)
combination of carbohydrates and lipids
combination of carbohydrates and proteins
molecule with the ability to bond with water; “water-loving”
molecule that does not have the ability to bond with water; “water-hating”
integral protein
protein integrated into the membrane structure that interacts extensively with the hydrocarbon chains of membrane lipids and often spans the membrane; these proteins can be removed only by the disruption of the membrane by detergents
peripheral protein
protein found at the surface of a plasma membrane either on its exterior or interior side; these proteins can be removed (washed off of the membrane) by a high-salt wash

Ribosomes - make protein for use by the organism
Cytoplasm - jelly-like goo on the inside of the cell
DNA - genetic material
Cytoskeleton - the internal framework of the cell
Cell membrane - outer boundary of the cell, some stuff can cross the cell membrane.

Prokaryotic Cells

Prokaryotes are very simple cells, probably first to inhabit the earth.
Prokaryotic cells do not contain a membrane bound nucleus.
Bacteria are prokaryotes.
DNA of bacteria is circular.

The word "prokaryote" means "before the nucleus"

Other features found in some bacteria:

Flagella - used for movement
Pilus - small hairlike structures used for attaching to other cells
Capsule - tough outer layer that protects bacteria, often associated with harmful bacteria

The structure, anatomy and morphology of mature seeds: model species in seed biology

Family (clade) Examples Description
Endospermic seeds
- Core Eudicots
- Rosid clade
muskmelon (Cucumis melo ) In the muskmelon seed the embryo is surrounded by a perisperm/endosperm envelope. Callose (ß-1,3-glucan) deposition in this envelope is responsible for the apoplastic semipermeability of muskmelon seeds. The perisperm/endosperm envelope is weakened prior to the completion of germination.
- Core Eudicots
- Rosid clade
fenugreek (Trigonella foenum-graecum)
crimson clover (Trifolium incarnatum)
lucerne (Medicago sativa)
Only some legume (Fabaceae) seeds are endospermic, most legume seeds are non-endospermic.
- Core Eudicots
- Rosid clade
castor bean (Ricinus communis) Castor bean seeds (Malpighiales) are a classical seed system to study endosperm reserve breakdown.
- Core Eudicots
- Rosid clade
garden cress (Lepidium sativum)
mouse-ear cress (Arabidopsis thaliana)
Only some Brassicaceae seeds are endospermic, most Brassicaceae seeds are non-endospermic. Mature seeds of Lepidium have 1-2 cell layers of endosperm, while Arabidopsis has a single endosperm cell layer. These two species exhibit, as tobacco, a two-step germination (distinct testa rupture and endosperm rupture). We found that Lepidium is a promising model system for endosperm weakening Müller et al. 2006).
- Core Eudicots
- Asterid clade
Cestroideae subgroup:
tobacco (Nicotiana tabacum)
other Nicotiana-species
petunia (Petunia hybrida)

Cestroideae subgroup of Solanaceae (tobacco, petunia):
Straight or slightly bent embryos and prismatic to subglobose seeds, two-step germination (distinct testa rupture and endosperm rupture), typically capsules as fruits.


All living organisms, regardless of their uniqueness, have certain biological, chemical, and physical characteristics in common. All, for example, are composed of basic units known as cells and of the same chemical substances, which, when analyzed, exhibit noteworthy similarities, even in such disparate organisms as bacteria and humans. Furthermore, since the action of any organism is determined by the manner in which its cells interact and since all cells interact in much the same way, the basic functioning of all organisms is also similar.

There is not only unity of basic living substance and functioning but also unity of origin of all living things. According to a theory proposed in 1855 by German pathologist Rudolf Virchow, “all living cells arise from pre-existing living cells.” That theory appears to be true for all living things at the present time under existing environmental conditions. If, however, life originated on Earth more than once in the past, the fact that all organisms have a sameness of basic structure, composition, and function would seem to indicate that only one original type succeeded.

A common origin of life would explain why in humans or bacteria—and in all forms of life in between—the same chemical substance, deoxyribonucleic acid ( DNA), in the form of genes accounts for the ability of all living matter to replicate itself exactly and to transmit genetic information from parent to offspring. Furthermore, the mechanisms for that transmittal follow a pattern that is the same in all organisms.

Whenever a change in a gene (a mutation) occurs, there is a change of some kind in the organism that contains the gene. It is this universal phenomenon that gives rise to the differences ( variations) in populations of organisms from which nature selects for survival those that are best able to cope with changing conditions in the environment.

Overview of Crosslinking and Protein Modification

A number of techniques for studying the structure and interaction of proteins, as well as for manipulating proteins for use in affinity purification or detection procedures, depend on methods for chemically crosslinking, modifying or labeling proteins.

Crosslinking is the process of chemically joining two or more molecules by a covalent bond. Modification involves attaching or cleaving chemical groups to alter the solubility or other properties of the original molecule. "Labeling" generally refers to any form of crosslinking or modification whose purpose is to attach a chemical group (e.g., a fluorescent molecule) to aid in detection of a molecule and is described in other articles.

The entire set of crosslinking and modification methods for use with proteins and other biomolecules in biological research is often called "bioconjugation" or "bioconjugate" technology. (Conjugation is a synonym for crosslinking.)

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Covalent modification and crosslinking of proteins depends on the availability of particular chemicals that are capable of reacting with the specific kinds of functional groups that exist in proteins. In addition, protein function and structure are either the direct focus of study or they must be preserved if a modified protein is to be useful in a technique. Therefore, the composition and structure of proteins, and the potential effects of modification reagents on protein structure and function, must be considered.

Proteins have four levels of structure. The sequence of its amino acids is the primary structure. This sequence is always written from the amino end (N-terminus) to the carboxyl end (C-terminus). Protein secondary structure refers to common repeating elements present in proteins. There are two basic components of secondary structure: the alpha helix and the beta-pleated sheet. Alpha helices are tight, corkscrew-shaped structures formed by single polypeptide chains. Beta-pleated sheets are either parallel or anti-parallel arrangements of polypeptide strands stabilized by hydrogen bonds between adjacent –NH and –CO groups. Parallel beta-sheets have adjacent strands that run in the same direction (i.e., N-termini next to each other), while anti-parallel beta sheets have adjacent strands that run in opposite directions (i.e., N-terminus of one strand arranged toward the C-terminus of adjacent strand). A beta-pleated sheet may contain two to five parallel or antiparallel strands.

Tertiary structure is the full three-dimensional, folded structure of the polypeptide chain and is dependent on the suite of spontaneous and thermodynamically stable interactions between the amino acid side chains. Disulfide bond patterns, as well as ionic and hydrophobic interactions greatly impact tertiary structure. Quaternary structure refers to the spatial arrangement of two or more polypeptide chains. This structure may be a monomer, dimer, trimer, etc. The polypeptide chains composing the quaternary structure of a protein may be identical (e.g., homodimer) or different (e.g., heterodimer).

The four levels of protein structure. The sequence of amino acids, represented by blue dots, joined by peptide bonds, comprise the primary structure. The properties of the constituent amino acids, in the context of the cellular environment, largely determine spontaneous formation of the higher-level structure that is essential for protein function.

Neurons Are Organized into Circuits

In complex multicellular animals, such as insects and mammals, various types of neurons form signaling circuits. In the simple type of circuit called a reflex arc, interneurons connect multiple sensory and motor neurons, allowing one sensory neuron to affect multiple motor neurons and one motor neuron to be affected by multiple sensory neurons in this way interneurons integrate and enhance reflexes. For example, the knee-jerk reflex in humans involves a complex reflex arc in which one muscle is stimulated to contract while another is inhibited from contracting (Figure 21-5). Such circuits allow an organism to respond to a sensory input by the coordinated action of sets of muscles that together achieve a single purpose. However, such simple nerve systems do not directly explain higher-order brain functions such as reasoning and computation.

Figure 21-5

The knee-jerk reflex arc in the human. Positioning and movement of the knee joint are accomplished by two muscles that have opposite actions: Contraction of the quadriceps muscle straightens the leg, whereas contraction of the biceps muscle bends the (more. )

The sensory and motor neurons of circuits such as the knee-jerk reflex are contained within the peripheral nervous system (Figure 21-6). These circuits send information to and receive information from the central nervous system (CNS), which comprises the brain and spinal cord and is composed mainly of interneurons. Highly specialized sensory receptor cells, which respond to specific environmental stimuli, send their outputs either directly to the brain (e.g., taste and odorant receptors) or to peripheral sensory neurons (e.g., pain and stretching receptors). The peripheral nervous system contains two broad classes of motor neurons. The somatic motor neurons stimulate voluntary muscles, such as those in the arms, legs, and neck the cell bodies of these neurons are located inside the central nervous system, in either the brain or the spinal cord. The autonomic motor neurons innervate glands, heart muscle, and smooth muscles not under conscious control, such as the muscles that surround the intestine and other organs of the gastrointestinal tract. The two classes of autonomic motor neurons, sympathetic and parasympathetic, generally have opposite effects: one class stimulates a muscle or gland, and the other inhibits it. Somatic sensory neurons, which convey information to the central nervous system, have their cell bodies clustered in ganglia, masses of nerve tissue that lie just outside the spinal cord. The cell bodies of the motor neurons of the autonomic nervous system also lie in ganglia. Each peripheral nerve is actually a bundle of axons some are parts of motor neurons others are parts of sensory neurons.

Figure 21-6

A highly schematic diagram of the vertebrate nervous system. The central nervous system (CNS) comprises the brain and spinal cord. It receives direct sensory input from the eyes, nose, tongue, and ears. The peripheral nervous system (PNS) comprises three (more. )

Having surveyed the general features of neuron structure, interactions, and simple circuits, let us turn to the mechanism by which a neuron generates and conducts electric impulses.


Some studies of plant anatomy use a systems approach, organized on the basis of the plant's activities, such as nutrient transport, flowering, pollination, embryogenesis or seed development. [4] Others are more classically [5] divided into the following structural categories:

About 300 BC Theophrastus wrote a number of plant treatises, only two of which survive, Enquiry into Plants (Περὶ φυτῶν ἱστορία), and On the Causes of Plants (Περὶ φυτῶν αἰτιῶν). He developed concepts of plant morphology and classification, which did not withstand the scientific scrutiny of the Renaissance.

A Swiss physician and botanist, Gaspard Bauhin, introduced binomial nomenclature into plant taxonomy. He published Pinax theatri botanici in 1596, which was the first to use this convention for naming of species. His criteria for classification included natural relationships, or 'affinities', which in many cases were structural.

It was in the late 1600s that plant anatomy became refined into a modern science. Italian doctor and microscopist, Marcello Malpighi, was one of the two founders of plant anatomy. In 1671 he published his Anatomia Plantarum, the first major advance in plant physiogamy since Aristotle. The other founder was the British doctor Nehemiah Grew. He published An Idea of a Philosophical History of Plants in 1672 and The Anatomy of Plants in 1682. Grew is credited with the recognition of plant cells, although he called them 'vesicles' and 'bladders'. He correctly identified and described the sexual organs of plants (flowers) and their parts. [6]

In the eighteenth century, Carl Linnaeus established taxonomy based on structure, and his early work was with plant anatomy. While the exact structural level which is to be considered to be scientifically valid for comparison and differentiation has changed with the growth of knowledge, the basic principles were established by Linnaeus. He published his master work, Species Plantarum in 1753.

In 1802, French botanist Charles-François Brisseau de Mirbel, published Traité d'anatomie et de physiologie végétale (Treatise on Plant Anatomy and Physiology) establishing the beginnings of the science of plant cytology.

In 1812, Johann Jacob Paul Moldenhawer published Beyträge zur Anatomie der Pflanzen, describing microscopic studies of plant tissues.

In 1813 a Swiss botanist, Augustin Pyrame de Candolle, published Théorie élémentaire de la botanique, in which he argued that plant anatomy, not physiology, ought to be the sole basis for plant classification. Using a scientific basis, he established structural criteria for defining and separating plant genera.

In 1830, Franz Meyen published Phytotomie, the first comprehensive review of plant anatomy.

In 1838 German botanist Matthias Jakob Schleiden, published Contributions to Phytogenesis, stating, "the lower plants all consist of one cell, while the higher plants are composed of (many) individual cells" thus confirming and continuing Mirbel's work.

A German-Polish botanist, Eduard Strasburger, described the mitotic process in plant cells and further demonstrated that new cell nuclei can only arise from the division of other pre-existing nuclei. His Studien über Protoplasma was published in 1876.

Gottlieb Haberlandt, a German botanist, studied plant physiology and classified plant tissue based upon function. On this basis, in 1884 he published Physiologische Pflanzenanatomie (Physiological Plant Anatomy) in which he described twelve types of tissue systems (absorptive, mechanical, photosynthetic, etc.).

British paleobotanists Dunkinfield Henry Scott and William Crawford Williamson described the structures of fossilized plants at the end of the nineteenth century. Scott's Studies in Fossil Botany was published in 1900.

Following Charles Darwin's Origin of Species a Canadian botanist, Edward Charles Jeffrey, who was studying the comparative anatomy and phylogeny of different vascular plant groups, applied the theory to plants using the form and structure of plants to establish a number of evolutionary lines. He published his The Anatomy of Woody Plants in 1917.

The growth of comparative plant anatomy was spearheaded by British botanist Agnes Arber. She published Water Plants: A Study of Aquatic Angiosperms in 1920, Monocotyledons: A Morphological Study in 1925, and The Gramineae: A Study of Cereal, Bamboo and Grass in 1934. [7]

Following World War II, Katherine Esau published, Plant Anatomy (1953), which became the definitive textbook on plant structure in North American universities and elsewhere, it was still in print as of 2006. [8] She followed up with her Anatomy of seed plants in 1960.


Two-component systems accomplish signal transduction through the phosphorylation of a response regulator (RR) by a histidine kinase (HK). Histidine kinases are typically homodimeric transmembrane proteins containing a histidine phosphotransfer domain and an ATP binding domain, though there are reported examples of histidine kinases in the atypical HWE and HisKA2 families that are not homodimers. [4] Response regulators may consist only of a receiver domain, but usually are multi-domain proteins with a receiver domain and at least one effector or output domain, often involved in DNA binding. [3] Upon detecting a particular change in the extracellular environment, the HK performs an autophosphorylation reaction, transferring a phosphoryl group from adenosine triphosphate (ATP) to a specific histidine residue. The cognate response regulator (RR) then catalyzes the transfer of the phosphoryl group to an aspartate residue on the response regulator's receiver domain. [5] [6] This typically triggers a conformational change that activates the RR's effector domain, which in turn produces the cellular response to the signal, usually by stimulating (or repressing) expression of target genes. [3]

Many HKs are bifunctional and possess phosphatase activity against their cognate response regulators, so that their signaling output reflects a balance between their kinase and phosphatase activities. Many response regulators also auto-dephosphorylate, [7] and the relatively labile phosphoaspartate can also be hydrolyzed non-enzymatically. [1] The overall level of phosphorylation of the response regulator ultimately controls its activity. [1] [8]

Phosphorelays Edit

Some histidine kinases are hybrids that contain an internal receiver domain. In these cases, a hybrid HK autophosphorylates and then transfers the phosphoryl group to its own internal receiver domain, rather than to a separate RR protein. The phosphoryl group is then shuttled to histidine phosphotransferase (HPT) and subsequently to a terminal RR, which can evoke the desired response. [9] [10] This system is called a phosphorelay. Almost 25% of bacterial HKs are of the hybrid type, as are the large majority of eukaryotic HKs. [3]

Two-component signal transduction systems enable bacteria to sense, respond, and adapt to a wide range of environments, stressors, and growth conditions. [11] These pathways have been adapted to respond to a wide variety of stimuli, including nutrients, cellular redox state, changes in osmolarity, quorum signals, antibiotics, temperature, chemoattractants, pH and more. [12] [13] The average number of two-component systems in a bacterial genome has been estimated as around 30, [14] or about 1–2% of a prokaryote's genome. [15] A few bacteria have none at all – typically endosymbionts and pathogens – and others contain over 200. [16] [17] All such systems must be closely regulated to prevent cross-talk, which is rare in vivo. [18]

In Escherichia coli, the osmoregulatory EnvZ/OmpR two-component system controls the differential expression of the outer membrane porin proteins OmpF and OmpC. [19] The KdpD sensor kinase proteins regulate the kdpFABC operon responsible for potassium transport in bacteria including E. coli and Clostridium acetobutylicum. [20] The N-terminal domain of this protein forms part of the cytoplasmic region of the protein, which may be the sensor domain responsible for sensing turgor pressure. [21]

Signal transducing histidine kinases are the key elements in two-component signal transduction systems. [22] [23] Examples of histidine kinases are EnvZ, which plays a central role in osmoregulation, [24] and CheA, which plays a central role in the chemotaxis system. [25] Histidine kinases usually have an N-terminal ligand-binding domain and a C-terminal kinase domain, but other domains may also be present. The kinase domain is responsible for the autophosphorylation of the histidine with ATP, the phosphotransfer from the kinase to an aspartate of the response regulator, and (with bifunctional enzymes) the phosphotransfer from aspartyl phosphate to water. [26] The kinase core has a unique fold, distinct from that of the Ser/Thr/Tyr kinase superfamily.

HKs can be roughly divided into two classes: orthodox and hybrid kinases. [27] [28] Most orthodox HKs, typified by the E. coli EnvZ protein, function as periplasmic membrane receptors and have a signal peptide and transmembrane segment(s) that separate the protein into a periplasmic N-terminal sensing domain and a highly conserved cytoplasmic C-terminal kinase core. Members of this family, however, have an integral membrane sensor domain. Not all orthodox kinases are membrane bound, e.g., the nitrogen regulatory kinase NtrB (GlnL) is a soluble cytoplasmic HK. [6] Hybrid kinases contain multiple phosphodonor and phosphoacceptor sites and use multi-step phospho-relay schemes instead of promoting a single phosphoryl transfer. In addition to the sensor domain and kinase core, they contain a CheY-like receiver domain and a His-containing phosphotransfer (HPt) domain.

The number of two-component systems present in a bacterial genome is highly correlated with genome size as well as ecological niche bacteria that occupy niches with frequent environmental fluctuations possess more histidine kinases and response regulators. [3] [29] New two-component systems may arise by gene duplication or by lateral gene transfer, and the relative rates of each process vary dramatically across bacterial species. [30] In most cases, response regulator genes are located in the same operon as their cognate histidine kinase [3] lateral gene transfers are more likely to preserve operon structure than gene duplications. [30]

Two-component systems are rare in eukaryotes. They appear in yeasts, filamentous fungi, and slime molds, and are relatively common in plants, but have been described as "conspicuously absent" from animals. [3] Two-component systems in eukaryotes likely originate from lateral gene transfer, often from endosymbiotic organelles, and are typically of the hybrid kinase phosphorelay type. [3] For example, in the yeast Candida albicans, genes found in the nuclear genome likely originated from endosymbiosis and remain targeted to the mitochondria. [31] Two-component systems are well-integrated into developmental signaling pathways in plants, but the genes probably originated from lateral gene transfer from chloroplasts. [3] An example is the chloroplast sensor kinase (CSK) gene in Arabidopsis thaliana, derived from chloroplasts but now integrated into the nuclear genome. CSK function provides a redox-based regulatory system that couples photosynthesis to chloroplast gene expression this observation has been described as a key prediction of the CoRR hypothesis, which aims to explain the retention of genes encoded by endosymbiotic organelles. [32] [33]

It is unclear why canonical two-component systems are rare in eukaryotes, with many similar functions having been taken over by signaling systems based on serine, threonine, or tyrosine kinases it has been speculated that the chemical instability of phosphoaspartate is responsible, and that increased stability is needed to transduce signals in the more complex eukaryotic cell. [3] Notably, cross-talk between signaling mechanisms is very common in eukaryotic signaling systems but rare in bacterial two-component systems. [34]

Because of their sequence similarity and operon structure, many two-component systems – particularly histidine kinases – are relatively easy to identify through bioinformatics analysis. (By contrast, eukaryotic kinases are typically easily identified, but they are not easily paired with their substrates.) [3] A database of prokaryotic two-component systems called P2CS has been compiled to document and classify known examples, and in some cases to make predictions about the cognates of "orphan" histidine kinase or response regulator proteins that are genetically unlinked to a partner. [35] [36]


The first class of biomolecules we will discuss are the carbohydrates. These molecules are comprised of the elements carbon (C), hydrogen (H), and oxygen (O). Commonly, these molecules are known as sugars. Carbohydrates can range in size from very small to very large. Like all the other biomolecules, carbohydrates are often built into long chains by stringing together smaller units. This works like adding beads to a bracelet to make it longer. The general term for a single unit or bead is a monomer . The term for a long string of monomers is a polymer .

Examples of carbohydrates include the sugars found in milk (lactose) and table sugar (sucrose). Depicted below is the structure of the monomer sugar glucose, a major source of energy for our body.

Carbohydrates have several functions in cells. They are an excellent source of energy for the many different activities going on in our cells. Some carbohydrates may have a structural function. For example, the material that makes plants stand tall and gives wood its tough properties is a polymer form of glucose known as cellulose . Other types of sugar polymers make up the stored forms of energy known as starch and glycogen . Starch is found in plant products such as potatoes and glycogen is found in animals. A short molecule of glycogen is shown below. You can manipulate the molecule yourself to take a good look.

Carbohydrates are essential for cells to communicate with each other. They also help cells adhere to each other and the material surrounding the cells in the body. The ability of the body to defend itself against invading microbes and the removal of foreign material from the body (such as the capture of dust and pollen by the mucus in our nose and throat) is also dependent on the properties of carbohydrates.

1.2 Themes and Concepts of Biology

By the end of this section, you will be able to do the following:

  • Identify and describe the properties of life
  • Describe the levels of organization among living things
  • Recognize and interpret a phylogenetic tree
  • List examples of different subdisciplines in biology

Biology is the science that studies life, but what exactly is life? This may sound like a silly question with an obvious response, but it is not always easy to define life. For example, a branch of biology called virology studies viruses, which exhibit some of the characteristics of living entities but lack others. Although viruses can attack living organisms, cause diseases, and even reproduce, they do not meet the criteria that biologists use to define life. Consequently, virologists are not biologists, strictly speaking. Similarly, some biologists study the early molecular evolution that gave rise to life. Since the events that preceded life are not biological events, these scientists are also excluded from biology in the strict sense of the term.

From its earliest beginnings, biology has wrestled with three questions: What are the shared properties that make something “alive”? Once we know something is alive, how do we find meaningful levels of organization in its structure? Finally, when faced with the remarkable diversity of life, how do we organize the different kinds of organisms so that we can better understand them? As scientists discover new organisms every day, biologists continue to seek answers to these and other questions.

Properties of Life

All living organisms share several key characteristics or functions: order, sensitivity or response to the environment, reproduction, adaptation, growth and development, regulation/homeostasis, energy processing, and evolution. When viewed together, these eight characteristics serve to define life.


Organisms are highly organized, coordinated structures that consist of one or more cells. Even very simple, single-celled organisms are remarkably complex: inside each cell, atoms comprise molecules. These in turn comprise cell organelles and other cellular inclusions. In multicellular organisms (Figure 1.10), similar cells form tissues. Tissues, in turn, collaborate to create organs (body structures with a distinct function). Organs work together to form organ systems.

Sensitivity or Response to Stimuli

Organisms respond to diverse stimuli. For example, plants can bend toward a source of light, climb on fences and walls, or respond to touch (Figure 1.11). Even tiny bacteria can move toward or away from chemicals (a process called chemotaxis) or light (phototaxis). Movement toward a stimulus is a positive response, while movement away from a stimulus is a negative response.

Link to Learning

Watch this video to see how plants respond to a stimulus—from opening to light, to wrapping a tendril around a branch, to capturing prey.


Single-celled organisms reproduce by first duplicating their DNA, and then dividing it equally as the cell prepares to divide to form two new cells. Multicellular organisms often produce specialized reproductive cells—gametes and oocyte and sperm cells. After fertilization (the fusion of an oocyte and a sperm cell), a new individual develops. When reproduction occurs, DNA containing genes are passed along to an organism’s offspring. These genes ensure that the offspring will belong to the same species and will have similar characteristics, such as size and shape.


All living organisms exhibit a “fit” to their environment. Biologists refer to this fit as adaptation, and it is a consequence of evolution by natural selection, which operates in every lineage of reproducing organisms. Examples of adaptations are diverse and unique, from heat-resistant Archaea that live in boiling hotsprings to the tongue length of a nectar-feeding moth that matches the size of the flower from which it feeds. Adaptations enhance the reproductive potential of the individuals exhibiting them, including their ability to survive to reproduce. Adaptations are not constant. As an environment changes, natural selection causes the characteristics of the individuals in a population to track those changes.

Growth and Development

Organisms grow and develop as a result of genes providing specific instructions that will direct cellular growth and development. This ensures that a species’ young (Figure 1.12) will grow up to exhibit many of the same characteristics as its parents.


Even the smallest organisms are complex and require multiple regulatory mechanisms to coordinate internal functions, respond to stimuli, and cope with environmental stresses. Two examples of internal functions regulated in an organism are nutrient transport and blood flow. Organs (groups of tissues working together) perform specific functions, such as carrying oxygen throughout the body, removing wastes, delivering nutrients to every cell, and cooling the body.

In order to function properly, cells require appropriate conditions such as proper temperature, pH, and appropriate concentration of diverse chemicals. These conditions may, however, change from one moment to the next. Organisms are able to maintain internal conditions within a narrow range almost constantly, despite environmental changes, through homeostasis (literally, “steady state”). For example, an organism needs to regulate body temperature through the thermoregulation process. Organisms that live in cold climates, such as the polar bear (Figure 1.13), have body structures that help them withstand low temperatures and conserve body heat. Structures that aid in this type of insulation include fur, feathers, blubber, and fat. In hot climates, organisms have methods (such as perspiration in humans or panting in dogs) that help them to shed excess body heat.

Energy Processing

All organisms use a source of energy for their metabolic activities. Some organisms capture energy from the sun and convert it into chemical energy in food. Others use chemical energy in molecules they take in as food (Figure 1.14).


The diversity of life on Earth is a result of mutations, or random changes in hereditary material over time. These mutations allow the possibility for organisms to adapt to a changing environment. An organism that evolves characteristics fit for the environment will have greater reproductive success, subject to the forces of natural selection.

Levels of Organization of Living Things

Living things are highly organized and structured, following a hierarchy that we can examine on a scale from small to large. The atom is the smallest and most fundamental unit of matter that retains the properties of an element. It consists of a nucleus surrounded by electrons. Atoms form molecules. A molecule is a chemical structure consisting of at least two atoms held together by one or more chemical bonds. Many molecules that are biologically important are macromolecules , large molecules that are typically formed by polymerization (a polymer is a large molecule that is made by combining smaller units called monomers, which are simpler than macromolecules). An example of a macromolecule is deoxyribonucleic acid (DNA) (Figure 1.15), which contains the instructions for the structure and functioning of all living organisms.

Link to Learning

Watch this video that animates the three-dimensional structure of the DNA molecule in Figure 1.15.

Some cells contain aggregates of macromolecules surrounded by membranes. We call these organelles . Organelles are small structures that exist within cells. Examples of organelles include mitochondria and chloroplasts, which carry out indispensable functions: mitochondria produce energy to power the cell, while chloroplasts enable green plants to utilize the energy in sunlight to make sugars. All living things are made of cells. The cell itself is the smallest fundamental unit of structure and function in living organisms. (This requirement is why scientists do not consider viruses living: they are not made of cells. To make new viruses, they have to invade and hijack the reproductive mechanism of a living cell. Only then can they obtain the materials they need to reproduce.) Some organisms consist of a single cell and others are multicellular. Scientists classify cells as prokaryotic or eukaryotic. Prokaryotes are single-celled or colonial organisms that do not have membrane-bound nuclei. In contrast, the cells of eukaryotes do have membrane-bound organelles and a membrane-bound nucleus.

In larger organisms, cells combine to make tissues , which are groups of similar cells carrying out similar or related functions. Organs are collections of tissues grouped together performing a common function. Organs are present not only in animals but also in plants. An organ system is a higher level of organization that consists of functionally related organs. Mammals have many organ systems. For instance, the circulatory system transports blood through the body and to and from the lungs. It includes organs such as the heart and blood vessels. Organisms are individual living entities. For example, each tree in a forest is an organism. Single-celled prokaryotes and single-celled eukaryotes are also organisms, which biologists typically call microorganisms.

Biologists collectively call all the individuals of a species living within a specific area a population . For example, a forest may include many pine trees, which represent the population of pine trees in this forest. Different populations may live in the same specific area. For example, the forest with the pine trees includes populations of flowering plants, insects, and microbial populations. A community is the sum of populations inhabiting a particular area. For instance, all of the trees, flowers, insects, and other populations in a forest form the forest’s community. The forest itself is an ecosystem. An ecosystem consists of all the living things in a particular area together with the abiotic, nonliving parts of that environment such as nitrogen in the soil or rain water. At the highest level of organization (Figure 1.16), the biosphere is the collection of all ecosystems, and it represents the zones of life on Earth. It includes land, water, and even the atmosphere to a certain extent.

Visual Connection

Which of the following statements is false?

  1. Tissues exist within organs which exist within organ systems.
  2. Communities exist within populations which exist within ecosystems.
  3. Organelles exist within cells which exist within tissues.
  4. Communities exist within ecosystems which exist in the biosphere.

The Diversity of Life

The fact that biology, as a science, has such a broad scope has to do with the tremendous diversity of life on earth. The source of this diversity is evolution , the process of gradual change in a population or species over time. Evolutionary biologists study the evolution of living things in everything from the microscopic world to ecosystems.

A phylogenetic tree (Figure 1.17) can summarize the evolution of various life forms on Earth. It is a diagram showing the evolutionary relationships among biological species based on similarities and differences in genetic or physical traits or both. Nodes and branches comprise a phylogenetic tree. The internal nodes represent ancestors and are points in evolution when, based on scientific evidence, researchers believe an ancestor has diverged to form two new species. The length of each branch is proportional to the time elapsed since the split.

Evolution Connection

Carl Woese and the Phylogenetic Tree

In the past, biologists grouped living organisms into five kingdoms: animals, plants, fungi, protists, and bacteria. They based the organizational scheme mainly on physical features, as opposed to physiology, biochemistry, or molecular biology, all of which modern systematics use. American microbiologist Carl Woese's pioneering work in the early 1970s has shown, however, that life on Earth has evolved along three lineages, now called domains—Bacteria, Archaea, and Eukarya. The first two are prokaryotic cells with microbes that lack membrane-enclosed nuclei and organelles. The third domain contains the eukaryotes and includes unicellular microorganisms (protists), together with the three remaining kingdoms (fungi, plants, and animals). Woese defined Archaea as a new domain, and this resulted in a new taxonomic tree (Figure 1.17). Many organisms belonging to the Archaea domain live under extreme conditions and are called extremophiles. To construct his tree, Woese used genetic relationships rather than similarities based on morphology (shape).

Woese constructed his tree from universally distributed comparative gene sequencing that are present in every organism, and conserved (meaning that these genes have remained essentially unchanged throughout evolution). Woese’s approach was revolutionary because comparing physical features are insufficient to differentiate between the prokaryotes that appear fairly similar in spite of their tremendous biochemical diversity and genetic variability (Figure 1.18). Comparing rRNA sequences provided Woese with a sensitive device that revealed the extensive variability of prokaryotes, and which justified separating the prokaryotes into two domains: bacteria and archaea.

Branches of Biological Study

The scope of biology is broad and therefore contains many branches and subdisciplines. Biologists may pursue one of those subdisciplines and work in a more focused field. For instance, molecular biology and biochemistry study biological processes at the molecular and chemical level, including interactions among molecules such as DNA, RNA, and proteins, as well as the way they are regulated. Microbiology , the study of microorganisms, is the study of the structure and function of single-celled organisms. It is quite a broad branch itself, and depending on the subject of study, there are also microbial physiologists, ecologists, and geneticists, among others.

Career Connection

Forensic Scientist

Forensic science is the application of science to answer questions related to the law. Biologists as well as chemists and biochemists can be forensic scientists. Forensic scientists provide scientific evidence for use in courts, and their job involves examining trace materials associated with crimes. Interest in forensic science has increased in the last few years, possibly because of popular television shows that feature forensic scientists on the job. Also, developing molecular techniques and establishing DNA databases have expanded the types of work that forensic scientists can do. Their job activities are primarily related to crimes against people such as murder, rape, and assault. Their work involves analyzing samples such as hair, blood, and other body fluids and also processing DNA (Figure 1.19) found in many different environments and materials. Forensic scientists also analyze other biological evidence left at crime scenes, such as insect larvae or pollen grains. Students who want to pursue careers in forensic science will most likely have to take chemistry and biology courses as well as some intensive math courses.

Another field of biological study, neurobiology , studies the biology of the nervous system, and although it is a branch of biology, it is also an interdisciplinary field of study known as neuroscience. Because of its interdisciplinary nature, this subdiscipline studies different nervous system functions using molecular, cellular, developmental, medical, and computational approaches.

Paleontology , another branch of biology, uses fossils to study life’s history (Figure 1.20). Zoology and botany are the study of animals and plants, respectively. Biologists can also specialize as biotechnologists, ecologists, or physiologists, to name just a few areas. This is just a small sample of the many fields that biologists can pursue.

Biology is the culmination of the achievements of the natural sciences from their inception to today. Excitingly, it is the cradle of emerging sciences, such as the biology of brain activity, genetic engineering of custom organisms, and the biology of evolution that uses the laboratory tools of molecular biology to retrace the earliest stages of life on Earth. A scan of news headlines—whether reporting on immunizations, a newly discovered species, sports doping, or a genetically-modified food—demonstrates the way biology is active in and important to our everyday world.

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    Watch the video: Prokaryotic vs. Eukaryotic Cells Updated (January 2022).