Showing posts with label physiology. Show all posts
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12:17 AM
Active transport is the process by which cells expend energy to move atoms or molecules across membranes, requiring the presence of a protein carrier, which is activated by ATP. Cotransport is active transport that uses a carrier that must simultaneously transport two substances in the same direction. Countertransport is active transport that employs a carrier that must transport two substances in opposite directions at the same time.

Biologists in nearly every field of study have discovered that one of the major methods by which organisms regulate their metabolisms is by controlling the movement of molecules into cells or into organelles such as the nucleus.

This regulation is possible because of the semipermeable nature of cellular membranes. The membranes of all living cells are fluid mosaic structures composed primarily of lipids and proteins. The lipid molecules are aliphatic, which means that their molecular structure exhibits both a hydrophilic (water-attracted) and a hydrophobic (water-repelling) portion.

These aliphatic molecules form a double layer: The hydrophilic heads are arranged opposite one another on the inner and outer surfaces, and the hydrophobic tails are aligned across from one another within the interior, sandwiched between the hydrophilic heads. The protein in the membrane is interspersed periodically throughout the lipid bilayer.


Some of the protein, referred to as peripheral protein, penetrates only one of the lipid layers. The integral protein, as the remaining protein is called, extends through both layers of lipid to interface with the environment on both the internal and external surfaces of the membrane. These integral proteins can serve as transport channels and carriers.

Cellular Energy

Transport across the membrane is accomplished by three different mechanisms: simple diffusion, facilitated diffusion, and active transport. The first two mechanisms are referred to as passive processes because they do not require the direct input of cellular energy, and they involve transport down a concentration gradient, that is, from the side with a higher concentration to the side with a lower concentration of the substance being transported.

In many instances, however, substances are transported across a membrane from the side with a low concentration to the side containing a greater concentration. This "uphill" movement across membranes is called active transport, and it requires the expenditure of cellular energy.

Cellular energy, produced by the biological oxidation of fuels such as carbohydrates, is stored as adenosine triphosphate (ATP). When this high-energy phosphate is hydrolyzed, the stored energy is released to drive cellular reactions such as active transport. The ATPase protein located in membranes belongs to one of the groups of enzymes which hydrolyze ATP.

The mechanism has not been completely deciphered, but it appears as though a protein carrier molecule binds with the substance to be transported at the surface on one side of the membrane. This binding occurs at a specific activated region on the carrier protein called the active site. After combining with the carrier, the substance is moved across the membrane and released at the surface on the other side of the membrane.

ATP is then hydrolyzed by an ATPase, and the energy released in this reaction prepares the protein carrier for attachment to another molecule to be transported by reactivating the active site. There is some question as to whether ATPase is a component of the carrier molecule or functions separately from it. Regardless of the spatial arrangement, the two molecules are intimately related in the active transport process.

Cotransport and Countertransport

There are two important modifications of the active transport process: cotransport and countertransport. Cotransport, or symport, involves a specialized protein molecule referred to as a symport carrier. Asymport carrier has two attachment sites. One site is for the attachment of the molecule to be transported, and the other is for the attachment of a second molecule, which can be referred to as the synergist.

Both the molecule to be transported and the synergist must be bound to the symport carrier before transport across the membrane can take place. The synergist is moved down a concentration gradient, and this downhill flow of the synergist drives the carrier to transport both molecules.

In order to keep the synergist moving down a concentration gradient when attached to the symport carrier, the synergist must be pumped back across the membrane. This movement of the synergist in the opposite direction is mediated by a protein carrier activated by the energy released from the hydrolysis of ATP by an ATPase.

Countertransport, or antiport, also utilizes a specialized carrier with two attachment sites. This antiport carrier binds with the molecule to be transported at one of the attachment sites, and a second molecule, which can be called the antagonist, binds at the other.

The carrier moves the molecule to be transported across the membrane while simultaneously moving the antagonist in the opposite direction. Again, both molecules must be attached to the antiport carrier before either can be transported, and the flow of the antagonist down a concentration gradient drives the transport by the carrier in both directions.

The antagonist is pumped back across the membrane by a protein carrier activated by the energy released from the hydrolytic action of an ATPase on ATP. This action maintains a concentration gradient favorable for transport when the antagonist is attached to the antiport carrier.

Transport in Action

The presence of these three active transport mechanisms has been well documented. Calcium, for example, has been shown to be pumped from the cell by a carrier protein activated by the hydrolysis of ATP. Sugars for energy and carbohydrate structure must be cotransported into the cell by a symport carrier that utilizes the sodium ion as a synergist. At least two countertransport ion pumps have been identified.

One pumps the potassium ion into the cell at the same time that it pumps the hydrogen ion out. The second pumps the potassium ion into the cell while the antagonist, sodium, is moved in the opposite direction. It is likely that numerous other active transport systems exist that have not yet been positively identified.

A protein carrier is one of the basic components of any active transport mechanism. Although no specific carrier molecule has yet been positively identified, there is ample indirect evidence to support the presence of such a protein. Much of this evidence comes from studies showing that active transport exhibits saturation kinetics.

This means that the transport of a specific ion will increase as the concentration increases, up to a certain point. At this point, further increases in concentration will have no effect on transport. These results strongly suggest that the ion is binding with another molecule in the membrane, such as a carrier protein, which is limited in concentration and becomes saturated.

Studies have also shown the transport of some substances to be competitively inhibited by the presence of another, very similar, substance. This indicates that both substances are competing for the same site on a membrane molecule, such as a protein carrier.

Role of Active Transport

The ability to accumulate substances against a concentration gradient is necessary for the normal function and survival of cells. There are numerous examples, however, of active transport being intimately involved in the regulation of some important biological processes. In the plant kingdom, sugar is produced by photosynthesis in the green leaves.

This sugar must be transported out of the leaves and into nonphotosynthetic tissues, such as roots or fruit, through specialized transport cells in the phloem. The loading of sugars into the phloem is dependent on an active cotransport mechanism.

Almost every field of life science is concerned with gene regulation. Genes are continually being induced (activated) or repressed (deactivated) as organisms develop and change from the time of their conception until their death.

Repression is usually caused by the presence of a protein molecule in the cell nucleus, but induction may very often be the result of metabolites being actively transported into the cell or nucleus. Hence, the active transport mechanisms may be a very important component of gene regulation.
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5:49 AM
Anaerobic Photosynthesis
Anaerobic Photosynthesis
Anaerobic photosynthesis, also known as anoxygenic photosynthesis, is the process by which certain bacteria use light energy to create organic compounds but do not produce oxygen. Anaerobes are those bacteria that cannot use oxygen to generate energy.

The photosynthetic process in all plants and algae, as well as in specific types of bacteria, involves the reduction of carbon dioxide to carbohydrate and the removal of electrons from water, resulting in the release of oxygen.

This process is known as oxygenic or aerobic photosynthesis. Water is oxidized by a multi-subunit protein located in the photosynthetic membrane. This is a molecular protein feature shared among more than 500,000 species of plants on earth.

While this is a common feature among nearly every form of plant life on earth, some photosynthetic bacteria can use light energy to extract electrons from molecules other than water. These bacteria are of ancient origin and are believed to have evolved before aerobic photosynthetic organisms.


These anaerobic photosynthetic organisms occur in the domain Bacteria. Anaerobic photosynthetic bacteria, also known as anoxygenic photosynthetic bacteria, differ from aerobic organisms in that each species of these bacteria has only one type of reaction center.

In some photosynthetic bacteria the reaction center involves the oxidation of water and the reduction of the aromatic molecule plastoquinone. In other species it involves the oxidation of plastocyanin and the reduction of ferredoxin protein.

Photosynthetic bacteria are typically aquatic microorganisms inhabiting marine and freshwater environments, including wet and muddy soils, stagnant ponds, sulfur springs, and still lakes. They are classified into five groups based on pigment composition, metabolic requirements, and membrane structure: green bacteria, purple sulfur bacteria, purple nonsulfur bacteria, heliobacteria, and halophilic archaebacteria.

Some of these organisms are strict anaerobes; that is, they can grow only in the complete absence of oxygen. They cannot use water as a substrate, and they do not produce oxygen during photosynthesis. Facultative anaerobes, on the other hand, can grow either in the presence or in the absence of oxygen.

Green bacteria include two families, the Chloroflexaceae and the Chlorobiaceae. The Chlorobiaceae are strict anaerobes that grow by utilizing sulfide, thiosulfate, or organic hydrogen as an electron source.

Anaerobic Photosynthesis Process
Anaerobic Photosynthesis Process

Chloroflexaceae are facultative aerobes which use reduced carbon compounds as electron donors. Purple sulfur bacteria uses an inorganic sulfur compound, such as hydrogen sulfide, as a photosynthetic electron donor.

Purple nonsulfur bacteria depend on the availability of simple organic compounds such as alcohols and acids as electron donors, but they can also use hydrogen gas. Purple sulfur bacteria must fix carbon dioxide to live, whereas nonsulfur bacteria can grow aerobically in the dark by respiration on an organic carbon source.

Heliobacteria are anaerobic photosynthetic bacteria that contain a special type of bacteriochlorophyll, BChl g, that works as both antenna and reaction center pigment. Halobacteria are very unusual. They cannot grow in low salt concentrations (or their cell walls collapse).

Typically, they are heterotrophs with an aerobic electron-transport chain, but they can also respire anaerobically, with nitrate or sulfur. In the absence of suitable electron acceptors they can ferment carbohydrates.

Halobacteria, when exposed to light in the absence of oxygen, can synthesize a purple membrane containing a single photosensitive protein called bacteriorhodopsin which, when illuminated, begins cyclic bleaching and regeneration, extruding protons from the cell. This light-stimulated proton pump operates without electron transport.

The mechanism by which halobacteria convert light is fundamentally different from that of higher organisms because there is no oxidation/reduction chemistry, and halobacteria cannot use carbon dioxide as their carbon source. As a result, some scientists do not consider halobacteria as being photosynthetic.

Process

The common features to both aerobic and anaerobic photosynthesis have been known since the mid-twentieth century:

Green plants:
CO2 + 2H2O + light → (CH2O) + O2 + H2O
Green sulfur bacteria:
CO2 + 2S + H2O + light → (CH2O) + 2S + H2O

In each case, inorganic carbon (CO2) is fixed into organic carbon (CH2O), the source of reductant is hydrogen in either water or hydrogen sulfide, and the chemical energy required for this activity is derived from light energy. The sulfur produced anaerobically is analogous to the oxygen produced by the oxygenic photosynthesis of green plants.

Photochemical processes in photosynthetic bacteria require three major components: an antenna of light-harvesting pigments, a reaction center within an intra-cytoplasmic membrane containing at least one bacteriochlorophyll, and an electron transport chain.

All photosynthetic bacteria can transform light energy into a trans membrane proton gradient used for the generation of adenosine triphosphate (ATP) and production of oxidase, but none of the anaerobic photosynthetic bacteria are capable of extracting electrons from water, so they do not evolve oxygen.

Many species can only survive in low-oxygen environments. To provide the necessary electrons for carbon dioxide reduction, anoxygenic photosynthetic bacteria must oxidize inorganic or organic molecules from their immediate environment.

Despite basic differences, the principles of energy transductions are the same in anaerobic and aerobic photsynthesis. Anaerobic photosynthetic bacteria depend on bacteriochlorophyll, a group of molecules similar to chlorophyll, that absorbs in the infrared spectrum between 700 and 1,000 nanometers. The antenna systems in these bacteria consist of bacteriochlorophyll and carotenoids, serving a reaction center where primary charge separation occurs.

Electron carriers include quinone and cytochrome bc complex. Electron transfer is coupled to the generation of electrochemical potential that drives phosphorylation by ATP synthase, and the energy required for the reduction of carbon dioxide is provided by ATP and dehydrogenase.
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3:07 AM
Angiosperm Cells and Tissues
Angiosperm Cells and Tissues
Some cell types and tissues which are not found in any other groups of plants occur in angiosperms (flowering plants).

Angiosperms are a group of plants with seeds that develop within an ovary and reproductive organs in flowers. They are commonly referred to as flowering plants and represent the most successful group of plants on earth, with approximately 235,000 species.

Various cell types and tissues, many of which are not found in any other groups of plants, occur in angiosperms. These cells and tissues perform varied functions, which are very efficient compared to their counterparts in other plants. These include dermal, vascular (xylem and phloem), and ground tissues (such as parenchyma, collenchyma, and sclerenchyma).

The growth of plants is carried on by a group of cells at their tips. These groups of cells are referred to as apical meristems, which are composed of initials and their most recent derivatives. The initials are the main source of body cells in plants,while the derivatives become any of the cells and tissues in the plant body.

The apical meristems of both the shoot and the root show continued cell division, with cells enlarging, elongating, and differentiating in regular, distinctly organized patterns. Apical meristems bring about the increase in the length of the stems and roots and are responsible in the formation of the primary plant body.

The shoot apical meristem may continually initiate the aerial components of the plant or may enter a state of periodic quiescence. In some plants, the shoot apical meristem transforms into a floral or inflorescence meristem that eventually terminates in a single flower or clusters of flowers, respectively.

The root apical meristem is enclosed by a thimble-shaped root cap that hastens the penetration of roots between soil particles.Unlike the shoot apical meristem, the root apical meristem forms no appendages. In fact, the site of lateral root initiation is far removed from it.

Shoot Apex

The shoot apical meristem is typically dome-shaped but flattened, and concave outlines also exist. The outline is not constant but changes in response to plastochron (the time interval between the initiation of one leaf primordium and the next). At least three models describe the shoot apical meristems. Although each of these is based on one or two unique criteria, they also have a few overlapping features.

Cell Lineage Analysis

This model holds that three clonally related layers of cells characterize the shoot apical meristem. These layers can be more than one cell layer thick. L1 is the outermost layer and gives rise to the epidermis, L2 is the middle layer and gives rise to the vascular tissues and cortex, and L3 is the inner most layer and gives rise to the pith.

This model was based on studies using periclinal chimeras (organs or parts of tissues of diverse genetic constitution), where one of the cell layers was genetically altered using drugs that inhibit separation of chromosomes.

Tunica-Corpus Concept

Tunica-Corpus Concept
Tunica-Corpus Concept
This model is based on microscopic analysis of constituent cells. It says that the shoot apical meristem is made up of two groups of cells. The tunica, a group of cells that form one or two stratified layers, undergoes anticlinal divisions only and gives rise to the epidermis.

Partly enclosed by the tunica is the corpus, a group of loosely arranged cells that divide in various planes and give rise to the vascular and ground tissues. The tunica maintains its individuality by surface growth, whereas the corpus adds bulk by increase in volume.

Cytohistological Zonation

This model recognizes various definable zones in the shoot apical meristem. Three zone boundaries are distinguished by cell size: staining quality, degree of vacuolation, and frequency of cell division.

The central (mother cell) zone represents a conspicuous group of enlarged and isodiametric cells that undergo infrequent cell division, possess prominent nuclei, and are often highly vacuolated. The flanking peripheral zone is derived from, and partly surrounds, the central zone. Cells of this zone are smaller, are mitotically active, and have dense cytoplasms.

They give rise to the epidermis, vascular tissues, and cortex. The rib zone is located at the base of the central and peripheral zones. This zone is directly formed from the central zone, produces longitudinal files of cells by periclinal divisions, and gives rise to the pith.

Root Apex

Root Apex
Root Apex
The organization of the root apical meristem is different from that of the shoot apical meristem. Root apical meristems are commonly interpreted as having either a close or open type of organization. In a close type of organization, the dermal, vascular, and ground tissues each have their own set of initials.

This organization shows a clear boundary between root cap and other tissues of the root apex. In an open type of organization, all of the root tissues share a group of initials, and therefore the boundary of the root cap is indistinguishable from the other tissues of the root apex.

Developmental Processes

The cells produced by apical meristems undergo several key developmental processes, which include growth, differentiation, and morphogenesis. Although each of these can be separated individually, they overlap in highly complex fashion.

Growth refers to the quantitative increase in a cell’s volume or mass due to enlargement and multiplication. Differentiation is the qualitative change in the form and function of organelles, cells, tissues, and organs, resulting in the establishment of new structures and functions.

From an anatomical point of view, cell differentiation is related to changes in cell size and shape, modifications of the wall, and changes in staining characteristics of nucleus or cytoplasm, as well as the degree of vacuolation and the ultimate loss of the protoplast in some cases. Morphogenesis is the visible manifestation of all of the changes, brought about by growth and differentiation, as expressed in the overall morphology of the plant.

Dermal Tissues

The primary plant body is composed of three basic tissues: dermal, vascular, and ground tissues. The dermal tissue (or epidermis) is made up of several cell types and is involved in a variety of functions, including retention and absorption of water and minerals, protection against herbivores, and control of gas exchange. Each of these functions is attributable to one or more of the unique features of the epidermis.

Most epidermal cells are flat and tightly packed, forming a single layer around stems, leaves, and other organs. The outer walls of epidermal cells are equipped with a waterproof layer made up of a fatty material called cutin. The tightly packed and cutinized epidermis protects the plants from desiccation by keeping moisture in.

Epidermal cells lack chloroplasts and are transparent. It is the underlying cells that give leaves and stems their green color. However, the vacuoles of some epidermal cells occasionally contain pigments and are responsible in the coloration of flowers and colored parts of variegated leaves.

Stomata are specialized structures that form part of the epidermis of leaves, stems, flowers, and fruits. They are involved in regulating the intake of carbon dioxide for photosynthesis as well as the release of oxygen. Trichomes are single-celled or multicellular out growths of epidermal cells that are involved in deterring herbivores and restricting transpiration.

Root hairs are also outgrowths of epidermal cells that are specialized for absorbing water and minerals from soil. They occur near the tip of the root and function to increase its absorptive surface area several-thousand fold.

Vascular Tissues

Vascular Tissues
Vascular Tissues
Vascular tissues are of two types: xylem and phloem. Xylem occurs throughout the plant body, and the type that differentiates directly from the apical meristem is called primary xylem. (Secondary xylem is formed from the vascular cambium.)

Primary xylem is formed as stems and roots elongate. The two kinds of conducting cells in xylem are tracheids and vessels, or vessel elements. Both are dead at maturity and have thick, lignified secondary cell walls. Tracheids are long, slender cells with tapered, overlapping ends.

They are the only water-conducting cells in most gymnosperms (an evolutionary line of plants that includes conifers).Water moves upward in roots and stems from tracheid to tracheid through thin areas in their cellwalls called pits. With only a few exceptions, all angiosperms contain vessel elements and tracheids.

Vessel elements are short, wide in diameter, and connected end to end. Their end walls are partly or wholly dissolved, forming long hollow vessels through which water moves. All these features of vessels enable them to transport water more rapidly than tracheids.

Phloem transports dissolved organic materials throughout the plant. The conducting cells of the phloem are called sieve elements, which are devoid of nuclei but otherwise have intact cytoplasm. They also have thin areas along their cell walls called sieve areas that are perforated. Solutes move from sieve element to sieve element through these pores.

Ground Tissues

The three types of ground tissue are parenchyma, collenchyma, and sclerenchyma. Parenchyma cells are the most abundant and versatile cells in plants. These cells are isodiametric, are alive at maturity, are highly vacuolated, and have a primary cell wall. Parenchyma functions as food-and-water-storage tissue as well as sites of metabolism in plants. Chlorenchyma cells are chloroplast-containing parenchyma specialized for photosynthesis.

Collenchyma cells are relatively long, with unevenly thickened primary walls. They support growing regions of shoots and are common in petioles, elongating stems and expanding leaves. Collenchyma cells are well adapted for support because their cell walls are able to stretch. They often form in strands or a cylinder just beneath the epidermis; such location maximizes support, as would a rod located in the center of a stemor petiole (leaf base).

Sclerenchyma cells are rigid; produce thick, non-stretchable secondary walls; and are usually dead atmaturity. They occur in, support, and strengthen mature regions of plants, including stems, roots, and leaves. There are two types of sclerenchyma cells: sclereids and fibers.

Sclereids are relatively short and variable in shape and usually occur in small groups. Fibers are long and slender and occur in strands or bundles. Sclereids are found in the roots, leaves, and stems.

They produce the gritty texture of pears and mostly make up the tough core of apples as well as the seed coats of peanuts and walnuts. Fibers are often associated with vascular tissues and, compared to sclereids, are typically elongated cells that vary in length from a few millimeters to more than half a meter long.
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11:03 PM
Angiosperm Life Cycle
Angiosperm Life Cycle

The word "angiosperm" comes from the Greek words for "vessel" and "seed" and translates roughly as "enclosed seed". In part, angiosperms (the flowering plants, phylumAnthophyta) are defined by the fact that their seeds are enclosed by an ovule. The life cycle of an angiospermis defined by the formation of the seed and its development to a full-grown plant which, in turn, produces seeds.

Angiosperms are vascular plants with flowers that produce seeds enclosed in an ovule—a fact that is recognized as the angiospermy condition.

Reproductive Flower Parts

In general, angiosperms have a floral axis with four floral parts, two of which are fertile. At the receptacle, or tip, of the axis there is an ovule-bearing leaf structure known as the carpel. The ovule or ovules can be found inside the pistil. Three portions compose the pistil: the ovary, the style, and the stigma, where the pollen usually germinates.


The mature ovule consists of the placenta, the integuments that are modified leaves that cover the entrance to the embryo sac, the micropyle, and the chalaza. These latter two parts of the ovule complement each other in their positions and functions.

While the micropyle receives and guides the pollen tube, the chalaza relates to the vascular supply of the ovule, nutrition, and support. The stamens, which are often composed of the filament and sporangia sacs that make up the anther, surround the pistil. Stamens carry the male gametes, and the pistil carries the female gamete needed for sexual reproduction.

It is believed that the great diversity and adaptability of the angiosperms is related to the presence of a unique reproductive cycle. This cycle consists of an alternation of generations and the production of a pair of spores on two types of sporophylls: microspores (which become male gametophytes) and megaspores (which become female gametophytes).

Male Gamete Development

The angiosperm reproductive cycle begins with the process of microsporogenesis, or microspore formation. The stamen consists of a filament and the anther, also known as the microsporangium. Inmost of the cases, the anther consists of four pollen sacs, or locules.

Within each locule, the archesporial cell develops through mitosis and extends as a row of cells throughout the entire length of the young anther. These mitotic cell divisions generate the anther wall, which is made up of several cell layers, the outermost of which transforms itself into the epidermis. The layer of cells belowthe epidermis is known as the endothecium.

During anther development, the endothecial cells acquire thickenings whose function is related to anther opening and pollen release. The innermost layer of the anther wall is the tapetum,whose primary function correlates with the nourishment of the young pollen and the deposition of the exine, a coating of the pollen grain.

As development proceeds, the sporogenous cells located below the tapetum transform into microsporocytes. These microsporocytes will undergo meiosis, and tetrads (units of four) of microspores will form.

Shortly after their formation, the tetrads separate into individual microspores. Each microspore is haploid, and often it will enlarge and separate from the tetrad, becoming sculptured by the deposition of sporopollenin and other substances that will turn into the ornamented surface of the pollen grain.

The second phase of pollen development is known asmicrogametogenesis. Themicrospore is the first cell of the gametophytic generation, the cell that generates themature pollen. The single-nucleus microspore develops into the male gametophyte before the pollen is released.

This developmental process occurs through two or three unequalmitotic divisions of the nucleus and subsequent cytokinesis (cell separation). The two daughter nuclei and cells differ in size and in form.

The larger cell represents the tube cell and nucleus,while the smaller cell represents the generative cell and nucleus. At maturity, the grain can be shred in two or three nucleate conditions. When the anther opens, the mature male gametophytes or pollen grains will be disseminated and ready for germination.

Female Gamete Development

The ovule (female sex organ) consists of two opposite ends: the micropyle, where the integuments come together, and a more distant end, where the ovular tissue is more massive. This part is also known as the chalaza, and it lies directly opposed to the micropyle.

The mature ovule is composed of three layers: the outer integument; the inner integument; and, underneath the integuments, the nucellus. During ovular development, one cell lying below the nucellar epidermis changes into a primary archesporial; this will divide to form the primary parietal cell and primary sporogenous cell.

The primary sporogenous cell functions as the megaspore mother cell, which divides meiotically, originating four haploid megaspores. In the majority of angiosperms, three of the megaspores will degenerate, and only the chalazal one will develop into the megagametophyte (embryo sac).

After the completion of the embryo sac stage, a series of cellular events occurs, ending with the formation of the mature embryo sac, ready for fertilization by the male gametes. The chalazal megaspore enlarges and undergoes threemitoses, giving rise to eight haploid cells. The mature megagametophyte consists of two groups of four cells located at both ends of the embryo sac.

The result is three antipodals at the chalazal end: the egg apparatus (consisting of the egg and two synergids at the micropylar end) and the polar nuclei. These two cells, present at both ends, usually fuse before pollination, and during fertilization they form the primary endosperm nucleus.

Pollination

The plant reproductive structures are now ready for the union of male and female gametes or fertilization, which eventually will produce a seed with a viable embryo and cotyledons. Before that step takes place, however, the pollen must be transferred from the anther to the stigma. Biotic agents (such as birds, insects, or mammals) or abiotic agents (such as wind or water) can accomplish this transfer process, known as pollination.

After landing on the stigma, pollen tubes will emerge through the grain apertures if the environ- ment is high in humidity. Successful germination of the pollen in the stigma requires nutrients. In most plants, growth of the pollen tube lasts between twelve and forty-eight hours, frompollen germination to fertilization.

Pollen germination starts with pollen-tube initiation, elongation, and penetration of the stigmatic tissue. During this period intense metabolic activity takes place, for the tube must synthesize membrane material and cell wall for growth and expansion. Simultaneously, at its tip the tube carries the vegetative cell nucleus, fol- lowed by the germinative cell.

Angiosperms have evolved complex breeding systems that ensure they will be pollinated by their own species. Today it is recognized that two pollination syndromes exist: self-pollination and cross-pollination. In self-breeding species, the pollen comes from the anther of the same flower.

In cross-pollination (or outcrossing) species, the pollen comes from the anthers of a different flower or even a different plant of the same species. In these plants, incompatibility in the stigma guarantees that only pollen from other flowers will germinate.

Fertilization

The union of one sperm with the egg is known as fertilization. However, several developmental processes in the vegetative and germinative cells prepare the two sperms for a process known as double fertilization. A mitotic division of the germinative cell generates the spermcells. This process that can take place on the growing pollen tube or inside the pollen grain.

In a growing pollen tube, the vegetative nucleus disintegrates and the sperm cells will take the lead and enter the embryo sac for successful fertilization. Usually, the interactions between the pollen grain and the pistil ensure that the sperm cells will often reach the micropyle of the ovule.

Once the spermreach themicropyle, the growth of other tubes stops. In the embryo sac (female gametophyte), four cells are located at themicropylar side.Of those four, the first pair that the spermcells will encounter are the synergids.

One of these is always bigger than the other and carries the filiform apparatus, a structure resembling hairs that degenerates after pollination and before fertilization. The synergids act as chemical attractants to the pollen tube, which penetrates the synergids via the filiform apparatus and then releases the two sperm cells.

One of the sperm cells will fuse with the egg, producing the zygote; the other sperm cell will fuse with the primary endosperm nucleus, generating the endosperm. The remaining cells of the female gametophyte are the antipodals; they usually degenerate after fertilization has taken place.

Seed and Fruit Formation

Once fertilization has occurred, the ovule will go through a series of metabolic steps ending with the formation of the seed and the fruit. The recently created zygote transforms into amulticellular and complex embryo that has two well-defined polar ends: the radicle, or primary root, and the embryonic apical meristem with the first leaves.

After successive mitosis, the mature endosperm usually grows close to the embryo and may provide nutrients needed for germination. The integuments will undergo further transformation, replication, and elongation and will become the seed coat—of variable texture, consistency, and colors, depending on the type of plant.

In general, after pollination or during fertilization, the ovary undergoes a series of physiological changes regulated by synchronized hormonal and genetic alterations that will modify the size of the parenchyma cells and its sugar and organic acids contents.

This process turns the ovary into fruit—in many cases familiar as the edible fruits familiar in human diets. The fruit provides nourishment for the seed until it ripens and drops to the ground, where the next stage in the life cycle begins.

Germination, Seedling Development, and Maturation

Seeds are released from the fruit in a large variety of ways that have evolved to ensure the survival of species. Whether ingested by mammals and passed through their feces to the ground, borne by wind on feathery "wings", or simply falling from rotting fruit that has abscissed and dropped from the plant, the seed must next undergo a process called germination, in which the embryo enclosed in the seed begins its growth. The embryo develops a hypocotyl (root axis) and a fleshy part known as the cotyledon; inmonocots there is one cotyledon, in dicots, two.

Germination requires certain conditions, such as the softening of the seed coat, moisture, and adequate warmth, to occur. During germination, the hypocotyl begins growing downward to become the root; the cotyledon(s) will develop into the shoot, stems, and leaves.

The process of germination results in the sprouting through the ground’s surface of the seedling, which will develop into the mature plant with flowers. The cycle then begins again.
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9:30 PM
Angiosperm Plant Formation
Angiosperm Plant Formation
Angiosperms are flowering plants. Their formation entails development from embryo to seed, through germination to seedling, and finally to mature plant.

The life cycle of angiosperms (flowering plants) involves an alternation of generations between a dominant sporophytic (spore-producing) phase and a reduced gametophytic (gamete-producing) phase. The first cell of the sporophyte is the fertilized egg, or zygote, which undergoes repeated divisions, growth, and differentiation to form an embryo enclosed in the ovule.

After fertilization, the ovule is transformed into the seed, which germinates into a seedling. The seedling becomes the adult plant; the plant produces flowers inwhich the sperm and egg—representing, respectively, the male and female gametophytic generations—are formed. Fertilization occurs, and seeds are produced to continue the life cycle.


Dicot Embryo Formation

In most angiosperms, embryo development, or embryogenesis, is initiated with a division of the fertilized egg into a small apical cell and a large basal cell, forming a two-celled proembryo. The apical cell generates the embryo proper, and the basal cell forms a filamentous suspensor that anchors the embryo.

Two weeds, Capsella bursa-pastoris (shepherd’s purse) and Arabidopsis thaliana (mouse ear cress or wall cress), both belonging to the Brassicaceae family, have attained prominence as textbook examples of embryogenesis in typical dicots (plants with two cotyledons, or seed leaves; a monocot has one seed leaf).

In these plants, the apical cell of the proembryo divides by two successive longitudinal walls to forma quadrant that is immediately partitioned by transverse walls into an octant, composed of an upper and lower tier of four cells each.

The fates of the two tiers are already fixed in the octant embryo, as the upper tier forms the shoot apex and much of the cotyledons. The lower tier, in addition to providing derivatives to the remaining part of cotyledons, generates the hypocotyl, the radicle, and the root apex.

However, the central region of the root cap, known as the columella, and the quiescent center of the root apical meristem are derived from the terminal cell of the suspensor closest to the embryo, known as the hypophysis. The apicobasal pattern of the future seedling plant is established in the octant embryo.

Aseries of divisions separating eight peripheral cells from a core of eight inner cells heralds histogenesis in the embryo. The result is the formation of a sixteen-celled, globular embryo, in which the peripheral cells form the protoderm (precursor cells of the embryonic epidermis), and the inner cells differentiate into the procambium and ground meristem (precursors of the vascular tissues and ground tissues, respectively) of the mature embryo. This initiates the formation of radial-pattern elements made up of concentric tissue layers in the basal part of the embryo.

Dicot Embryo Formation
Dicot Embryo Formation
The globular stage of the embryo is completed by approximately three additional rounds of divisions, mostly in the inner core of cells. The suspens or attains its genetically permissible number of six to nine cells by this stage. Gradually the cells begin to lose connection from one another and from the embryo and disintegrate.

Emerging from the globular stage, the embryo expands laterally by cell divisions to formthe cotyledons and becomes heart-shaped. The heart-shaped stage is followed by the torpedo-shaped stage, in which elongation of the cotyledons and hypocotyl, as well as extension of the vascular tissues, occurs.

The basic body plan of a shoot-root axis becomes unmistakably clear at this stage, with the establishment of the shoot apical meristem in the depression between the cotyledons and the organization of a root apex by incorporation of derivatives of the hypophysis at the opposite end of the embryo.

During further growth, the cotyledons bend toward the hypocotyl (bent cotyledon or walking-stick shaped stage), and the embryo is phased into the mature stage. A mature embryo of Arabidopsis has fifteen thousand to twenty thousand cells and, under favorable conditions of growth, develops in about nine days fromthe time of fertilization to the mature embryo stage.

Sensitive genetic screens have led to the isolation of Arabidopsis mutants defective in apicobasal and radial patterning of embryos. Characterization of the mutant genes and their protein products has unraveled to some extent the molecular components of the embryonic pattern-forming system in this plant.

Monocot Embryo Formation

The early divisions of the zygote in monocots follow the same pattern as in dicots. However, in the Poaceae (grasses) family, which includes wheat and the other cereals, the sequence and orientation of later divisions in the proembryo are irregular, resulting in highly complex mature embryos. The main feature of the cereal embryo is the development of an absorptive organ known as the scutellum (considered equivalent to the single cotyledon).

Other organs of the embryo for which there are no counterparts in the dicot embryo are a sheath like tissue covering the root (coleorhiza), a tissue that covers the shoot (coleoptile), and an internode known as the mesocotyl. On one side of the coleorhiza there is also a small, flaplike out growth called the epiblast.

Embryo Maturation to Seed

Embryo Maturation to Seed
Embryo Maturation to Seed
As the embryo matures, the ovule progressively desiccates to become the seed enclosed within the ovary. Concomitantly, the integuments of the ovule harden to form the protective seed coat. Within the ovule itself, the primary endosperm nucleus formed after double fertilization begins to divide, ahead of the zygote, to produce the endosperm charged with nutrient substances. In seeds ofmany plants, including Arabidopsis, Capsella, bean, and pea, the endosperm is utilized by the developing embryo.

In other plants, especially the cereals, the bulk of the seed (grain) is made up of the endosperm surrounding the small embryo. The mature embryo enclosed in the seed consists of an axis bearing the radicle (embryonic root) at one end and the plumule (the embryonic shoot consisting of the shoot apex and one or two leaves) at the other end, and one (in monocots) or two (in dicots) cotyledons.

The part of the embryo axis above the point of attachment of the cotyledon(s) is known as the epicotyl, whereas the part below the attachment point connecting to the radicle is called the hypocotyl.

Seed Germination

The dry seed enclosing the mature embryo may not germinate immediately; if it does not, it enters a period of quiescence or dormancy. Quiescent seeds germinate when provided with the appropriate conditions for growth, such as water, a favorable temperature, and the normal composition of the atmosphere.

Dormant seeds germinate only when some additional hormonal, environmental, metabolic, or physical conditions are met. In almost all seeds, the first part of the embryo to emerge during germination is the radicle. It forces it way through the soil and forms the primary root of the seedling. However, the manner in which the shoot emerges and develops varies considerably in different seeds.

In the epigeous type of germination (for example, in beans), emergence of the radicle is followed by the elongation of the hypocotyl, which arches above the soil surface as a hook. As the hook straightens, it pulls out the cotyledons and plumule above the soil surface. In the hypogeous type of germination (in peas, for example) the cotyledons enclosed within the seed coat remain in the soil during germination.

It is the epicotyl that arches above the soil surface, and as the hook straightens out, it carries the plumule along with it to the surface of the soil. In the monocot, such as the onion, after emergence of the radicle the single cotyledon arches above-ground and subsequently straightens.

Members of the Poaceae display a type of germination in which, following the outgrowth of the radicle, the coleoptile enclosing the plumule grows out of the grain and appears above the soil. The seedling leaves force theirway, breaking the coleoptile, and appear outside as the first photosynthetic organs.

The growth of the coleoptile during germination of grains is facilitated by the elongation of the mesocotyl. These various types of germination ensure an efficient use of food materials stored in the embryo or in the endosperm for the growth of the seedling until it becomes autotrophic.

Embryo to Adult Plant

Although the question as to whether the seedling will become a gigantic tree or a small, herbaceous plant is determined by its genetic blueprint, certain common postgermination growth and developmental episodesmark the development of the seedling into an adult plant.

In dicots, continued growth of the primary root produces an extensively branched root system consisting of secondary roots or lateral roots. In monocots, the primary root disintegrates shortly after it is formed, and so the root system is constituted of numerous adventitious roots which arise from the base of the stem.

Although the cotyledons retain their photosynthetic capacity for some time after germination of the seed, the seedling becomes completely autotrophic as the shoot apex produces new leaves and branches arise in the axils of leaves.

These activities are coordinated by the division of cells in the root and shoot apical meristems and the differentiation of cells into specialized tissues and organs. The shoot and root apical meristems, considered analogous to the stem cells of animals, remain active throughout the life of the plant and, hence, are known as indeterminate meristems.
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Biochemical Coevolution in Angiosperms
Biochemical Coevolution in Angiosperms
Flowering plants, or angiosperms, produce many compounds that are not directly related to growth and development. These secondary metabolites arise from primary metabolic pathways and act as antiherbivory mechanisms, allelochemicals, or attractants.

Secondary metabolites are biochemicals produced by plants in response to selection pressures. These pressures may be from herbivory, competition, or the need for pollination.

As plants produce compounds to enhance their survival, predators, competitors, and pollinators react and evolve means of adjusting to the plant’s efforts. Chemically simple secondary metabolites may be widespread throughout angiosperm (flowering plant) families, whereas more complex chemicals are often restricted to a single species.

Secondary metabolites are often under high selection pressures, causing individual compounds to have very limited distributions and making them useful in determining the evolutionary relationships between taxonomic groups. Presence of secondary compounds influences the activities of organisms interacting with the plants and, over long periods of time, influences evolution of those species.

Antiherbivory Mechanisms

Antiherbivory chemicals may have a wide range of effects on herbivores (plant-eating animals). Many compounds merely deter grazing. Crystals produced from calcium oxalate (raphides) may be ejected from the vacuoles of cells along with proteinaceous toxins, causing tissue swelling in the mouth of an offending herbivore.

Many monocotyledonous plant families, such as Liliaceae, Heliconiaceae, Rubiaceae, and Arecaceae, produce this type of antiforaging device. This type of defense is especially notable in young tissues that have not developed the toughness found inmature leaves as a herbivory deterrent. Red oaks produce tannins in response to gypsy moth attacks, reducing further herbivory.

Continued feeding on plants containing tannins would lead to slow starvation of a herbivore, as its digestive system could not absorb proteins. The same tannins are not deterrents to squirrels. Squirrels harvest acorns and bury them for later consumption, providing a food source for the squirrel and a dispersal mechanism for the acorn.

Beavers provide another example of the interaction between plant and animal evolution. Some of the beaver’s preferred foods include species that are unpalatable or toxic to other mammals, such as bracken fern, nettles, thistles, and skunk cabbage. This gives the beaver a largely uncontested food source that may involve a metabolic "cost" to the animal.

Other antiherbivory chemicals result in effects more severe than mere deterrence of feeding. Alkaloids such as caffeine, nicotine, and strychnine are potent anti herbivory mechanisms, causing convulsions, comas, and even death in herbivores. These effects may not occur in all herbivores.

Strychnine, for example, is produced by the fruit of some plants that may be eaten by birds without ill effects, but in mammals the same fruit causes failure of the central nervous system and induces seizures.The plant reduces herbivory by mammals, and the seeds get dispersed by birds that are able to detoxify the strychnine.

Grains and seed crops, such as wheat and peanuts, which are particularly attractive to animal and insect herbivores, often produce cyanogenic glycosides that release hydrogen cyanide as the tissues are digested. This compound inhibits cellular respiration, thus killing the herbivore. In each case members of a plant population are consumed by the herbivore, but future generations are spared by the loss.

Allelochemicals

Allelochemicals
Allelochemicals
Allelochemicals are compounds produced by an organism that interfere with the growth or development of another organism.Many phenolic acids act as allelo chemicals, inhibiting root growth of competing species. Many grains are known to release ferulic acid and caffeic acid into the soil, thus inhibiting the germination of weed species.

Phenolics may also act as antifungal compounds, increasing in concentration with fungal infection, thus protecting the plant from further attack. Phenolic compounds produced in tobacco and tomato leaves reduce the growth of these plants’ natural predator, tobacco hornworm, without affecting the growth or activity of the hornworm’s natural predator.

Allelochemicals produced in response to injury by herbivores may also attract predators of the herbivore.Wastes from many species of caterpillars induce the release of terpenoids from green leaves that attract parasitoid insects.

Production of specific combinations of volatiles on the part of the plant signals the predator, which will then reduce further herbivory. The plants have evolved the signal in response to herbivory, and the predators have evolved the ability to detect the signal indicating the location of their host.

Lectins are widely distributed carbohydrate-binding proteins, most commonly found in the Leguminoseae (legume) family. When found in the seeds, these compounds act as broad-spectrum insecticides, whereas in the roots of legumes they maintain bacterial relationships in nitrogen fixing nodules, providing the plant with a source of nitrogen unavailable to plants not producing nodules.

Attractants

Attractants
Attractants
Terpenoids and aliphatic compounds are often the components of essential oils of plants. The volatile nature of these compounds produces a distinctive odor that attracts pollinators. Composition of the volatile compounds often closely matches the natural pheromones produced by the pollinator, mimicking the chemical scent of a female insect in an attempt to attract male pollinators.

Pheromone mimicry is found primarily in members of the Orchidaceae, which are often dependent on single species of wasp for pollination. Other plants may mimic the odor of food. The smell of rotting flesh, attractive to flies, is produced via ammonia and alkylamines, such as cadaverine and putrescine. Methylesters may attract moth pollinators by mimicking the sweet smell of fruit.

Flavonoids often provide color to fruits and flowers and act as visual cues for pollination. Reds, blues, or yellows in varying patterns stand out against a background of green leaves, helping pollinators locate the flower.

Species may have minor chemical differences in their flavonoids that allow for the determination of identity, hybridization between species, and possible coevolution with pollinators. For example, tropical flowers tend to have a more intense red color from anthocyanins than do temperate flowers.

This difference correlates with differences in pollinator preferences, indicating a role by natural selection. Birds, such as hummingbirds, prefer red to yellow, whereas bees are not able to discern reds but are attracted to yellows. Carotenoids, such as xanthophyll and beta-carotene, give fruits and flowers distinctive yellow and orange colors.

Color patterns are also important in attracting pollinators. Butterflies are attracted to red/yellow color patterns. Flavonoid compounds not only impart color but also may modify color patterns by absorbing ultraviolet (UV) light.

Bees are capable of seeing in the UV range, so the presence of flavonoids may alter the bees’ perception of the flower. The patterns may also create cues as to the location of nectaries within the flower, guiding the pollinator to its reward.
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8:17 PM
Bioluminesence
Bioluminesence
Bioluminesence is the production of light by living organisms, including algae and phytoplankton in the oceans and fungi on land.

Bioluminescence is a specific form of chemiluminescence in which the chemical energy that is produced in a chemical reaction is converted into radiant energy. In bioluminescence the reaction originates in a wide variety of living organisms, including a small number of plants. It should not be confused with fluorescence or phosphorescence, both of which do not involve a chemical reaction.

In either of the former cases the energy from a source of light, not from a chemical reaction, is basically absorbed and then re-emitted in some form of another photon. The chemical reactions that lead to bioluminescence release energy in the form of light.

Unlike the light bulb, in which electrical energy is converted into light, with some of this energy lost in the form of heat, a bioluminescent reaction is 100 percent efficient and converts all the emitted energy into light. Because there is no heat released, bioluminescence is also known as "cold light."

Species and Habitats

Bioluminescence is primarily marine in nature and is the only source of light in the deep ocean, which is the largest habitable biome of the earth. The phenomenon rarely occurs in any source of fresh water. Bioluminescent organisms include ctenophores, annelid worms, mollusks, insects, and fish.

Species and Habitats
Species and Habitats
The most common manifestation of this phenomenon on land is seen as a glowing fungus on wood or in the few families of luminous insects. This property can be used as ameans of species recognition in the darkness as well as for courtship, preying, and mating.

There are several bioluminescent fungi that are not marine in nature, occurring primarily in the tropics. These fungi appear in different colors. The most common is Panellus stiptucus,which is a small decay fungus that is mostly restricted to North America. The jack-o’-lantern mushroom (Omphalotus olearius) glows brightly, especially when fresh. A few Armillaria species are also reported to glow mildly. No luminous tree or plant is known, however.

Mechanisms of Bioluminescence

Bioluminescence occurs only when two different species are in contact and, almost exclusively, when oxygen is present. The two species are luciferin, which produces the light, and luciferase, a protein that triggers and catalyzes the reaction. The mechanism involves the loss of two electrons, also known as oxidation, by luciferin, a process achieved only through the intervention of luciferase to yield oxyluciferin.

Occasionally luciferin, luciferase, and a cofactor such as oxygen are bound together in a single moiety called photoprotein, which leads to light formation upon contact with a positively charged species, such as the calcium cation. The mechanism appears to involve a peroxide decomposition with free radical intervention.

Dinoflagellates

Dinoflagellates known as Pyrrhophyta, or fire plants, are the most common sources of bioluminescence at the surface of the ocean. They are a group of marine algae that produce light upon mechanical, chemical, or temperature changes. The phenomenon was first observed in the genus Noctiluca in the nineteenth century and has since been observed to occur within other species.

Generally, three types of stimuli can cause bioluminescence in dinoflagellates: mechanical, chemical, and temperature stimulation.

Dinoflagellates
Dinoflagellates
Mechanical forms of stimulation, such as the stirring of water from a moving boat, a swimming fish, or a breaking wave, are prevalent in many Pyrrhophyta. The light appears to serve as a "burglar alarm" against grazing predators, which are then being seen through the flash by a larger second predator.

For example, as a copepod approaches the dinoflagellate, agitation of the seawater stimulates light flashes which a small fish, the secondary predator, uses to pinpoint the position of the copepod and eventually consume it. It appears that the mechanical stimulation deforms the cell membrane to create a short flash as little as one one-hundredth of a second.

Dinoflagellate luciferin is thought to derive from the similarly structured chlorophyll, which is found in most plants. The molecule is protected from luciferase at slightly basic medium by a luciferin-binding protein. However, once the acidity increases, the free luciferin reacts, and light is emitted.

The light produced by a single dinoflagellate is only six to eight photons in energy, and the flashing may last only one-tenth of a second. Larger organisms, such as jellyfish, provide flashes that may last up to tens of seconds. Temperature lowering in some dinoflagellate species also creates bioluminescence.

Purpose and Applications

The disappearance of the flash, once oxygen is consumed, has suggested that the bioluminescent reaction was originally used to remove toxic oxygen from primitive types of bacteria that developed at a time when oxygen was not available.

Bioluminescence has also played a crucial role in the direct studies of several cellular and biochemical processes, such as in the formation of ultimate carcinogens from benzoapyrene. The phenomenon has served scientists in many ways.

Calcium levels are monitored via the jellyfish biochemical system, adenosine triphosphate (ATP) measurements are achieved through the firefly, and the gene activity of organisms can be detected by splicing known bioluminescent proteins.
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1:51 AM
Bulbs and rhizomes are modified stems, stem bases, or other underground organs used by plants for food (or energy) storage and in asexual reproduction.

Plants reproduce both sexually and asexually. Although sexual reproduction is part of the typical life cycle of plants, for a variety of reasons a plant may reproduce asexually. Exact duplicates of a plant, called clones, are formed by asexual reproduction.

Asexual Reproduction

Asexual reproduction involves the production of offspring through the formation of propagules by mitosis (the process of nuclear cell division). Because genetic recombination does not occur in mitosis, the offspring are genetically identical to the parent plant.

Asexual reproduction does not occur in all plants; some reproduce asexually only when humans intervene. Asexual reproduction occurs when a single plant produces a vegetative propagule that develops into a separate free-living plant. Many of the propagules that support asexual reproduction are actually highly modified branches. Others are modified roots.


In rare instances, the tissues of leaves may be modified by nature to support asexual reproduction. The propagules of asexual reproduction vary enormously. They are often found in catalogs describing “bulbs,” but technically they include true bulbs, corms, stolons, tubers, rhizomes, turions, pseudobulbs, and fleshy roots.

True Bulbs

Bulbs, corms, stolons, tubers, rhizomes, and turions are all modified stems. Bulbs are modified stem bases that develop underground. The stem is shortened and thickened to produce a mass of tissue shaped like a coin or like a child’s toy top. Scale like leaves with thickened bases are attached to the base of the bulb.

Starch is stored in the thickened bases, a food supply that allows the bulb to survive through a dormant season and to produce adventitious roots. Roots are often absent when the bulb is dormant. The starch can also support a period of rapid stem and leaf growth in the growing season and may support the flowering and fruiting of the plant.

In tunicate bulbs, a cloak of dried leaves surrounds the outside of the bulb. These dried leaves provide a barrier to desiccation and allow the tunicate bulbs to be stored above ground for weeks or even months.

Onions (Allium cepa) and daffodils (Narcissus) are examples of tunicate bulbs. Other bulbs have no cloak and usually have shorter, less cylindrical leaves. These scaly bulbs dry quickly when kept above ground and usually develop flowers only after a more normal, aerial branch system forms. Lilies (Lilium) have scaly bulbs.

Stems of both tunicate and scaly bulbs can branch. Below ground, branches appear at first as miniature bulbs (bulbils or bulblets). Bulblets take their energy from the parent bulb but eventually produce aerial stems or leaves and can be separated from the parent.

Profuse branching can be stimulated by wounding the stem of the parent bulb. All the bulbs produced by this technique are clones, identical to the parent in their genetic makeup and physical characteristics.

Corms and Tubers

Corms are similar to bulbs in many ways. They have a disk-shaped or top-shaped stem mass that is shorter and wider than most typical stems. They are often cloaked in a tunic of dried leaves that are thinner and smaller than those on bulbs.

Corms do not store significant amounts of starch; it is instead stored primarily in the basal plate of the stem. Branches of the corm stem produce new, miniature corms (cormels). Wounding the parent stem stimulates greater branching. Gladiolus and crocus are two common garden plants that produce corms.

Tubers are thick, starchy stems that form usually at the tip of a stolon, runner, or tiller. Tubers may form on the soil surface or below ground. A familiar example is the white or Irish potato (Solanum tuberosum). The leaves on most tubers are much smaller than the leaves on other parts of the stem, but above each leaf on the tuber is a well developed axillary bud, commonly called an eye.

The axillary bud has the potential to elongate, forming a complete and fully developed branch. If the stolon connecting the tuber to the parent plant dies, the branch from the eye of the tuber becomes an independent clone of the parent plant.

Most tubers contain many eyes. If the tuber is cut into smaller pieces, each containing an eye, each piece develops a rhizome, which in turn develops a newtuber. By this technique, a significant increase in the number of plants can be obtained.

The cut pieces of tuber are initially prone to decay, but after they have dried for a few days, they heal over with a layer of callus which protects them like a skin. Cutting tubers into small “seed” pieces is a common method for propagating tuber-forming species.

Rhizomes, Stolons, and Fleshy Roots

Rhizomes are specialized, underground stems. Unlike most areal stems, the rhizomes are normally oriented horizontally. Just as pieces of tuber can provide the tissue and energy source for the formation of a new plant, so too can pieces of a rhizome.

Many ferns and fern allies propagate naturally by rhizomes. Large stands of these plants can form from a single individual as the rhizomes grow and branch. Eventually, older pieces of the rhizome die, leaving a population of individuals that all have identical genetic characteristics.

Stolons are long, thin, horizontal stems, also called runners or tillers, which grow along the surface of the ground. When the stolon has grown far enough from the parent plant, the growth pattern changes, and a crown, or tuber, forms. A crown is a compressed stem mass with leaves arranged close to one another, also called a rosette. Within the crown, roots format the points of attachment of the leaves to the stems.

If the stolon is broken or dies, the crown becomes an independent clone of the parent plant. In this way, a large number of off-spring can be produced from a single plant. This is a mechanism of reproduction of the strawberry (Fragaria) and is also a common reproductive mechanism for grasses, including crab grass (Digitaria sanguinalis) and quack grass (Agropyron repens).

Fleshy roots store energy that can be useful for asexual propagation. Most require at least a small amount of stem tissue to support cell growth and differentiation. The true yam (Dioscorea) is a tuber, but the sweet potato (Ipomoea batatas) is a fleshy root which can easily be propagated asexually. Many buttercups (Ranunculus) are also propagated by breaking up the clumps of their fleshy roots.
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11:45 PM

Alternative forms of photosynthesis are used by specific types of plants, called C4 and CAM plants, to alleviate problems of photorespiration and excess water loss.

Photosynthesis is the physiological process whereby plants use the sun’s radiant energy to produce organic molecules. The backbone of all such organic compounds is a skeleton composed of carbon atoms. Plants use carbon dioxide from the atmosphere as their carbon source.

The overwhelming majority of plants use a single chemical reaction to attach carbon dioxide from the atmosphere onto an organic compound, a process referred to as carbon fixation. This process takes place inside specialized structures within the cells of green plants known as chloroplasts.


The enzyme that catalyzes this fixation is ribulose bisphosphate carboxylase (Rubisco), and the first stable organic product is a three-carbon molecule. This three-carbon compound is involved in the biochemical pathway known as the Calvin cycle. Plants using carbon fixation are referred to as C3 plants because the first product made with carbon dioxide is a three-carbon molecule.

C4 Photosynthesis

For many years scientists thought that the only way photosynthesis occurred was through C3 photosynthesis. In the early 1960’s, however, researchers studying the sugarcane plant discovered a biochemical pathway that involved incorporation of carbon dioxide into organic products at two different stages.

First, carbon dioxide from the atmosphere enters the sugarcane leaf, and fixation is accomplished by the enzyme phosphoenolpyruvate carboxylase (PEP carboxylase). This step takes place within the cytoplasm, not inside the chloroplasts. The first stable product is a four-carbon organic compound that is an acid, usually malate. Sugarcane and other plants with this photosynthetic pathway are known as C plants.

In C4 plants, this photosynthetic pathway is tied to a unique leaf anatomy known as Kranz anatomy. This term refers to the fact that in C4 plants the cells that surround the water- and carbohydrate conducting system (known as the vascular system) are packed very tightly together and are called bundle sheath cells.

Surrounding the bundle sheath is a densely packed layer of mesophyll cells. The densely packed mesophyll cells are in contact with air spaces in the leaf, and because of their dense packing they keep the bundle sheath cells from contact with air. This Kranz anatomy plays a major role in C4 photosynthesis.

In C4 plants the initial fixation of carbon dioxide from the atmosphere takes place in the densely packed mesophyll cells. After the carbon dioxide is fixed into a four-carbon organic acid, the malate is transferred through tiny tubes from these cells to the specialized bundle sheath cells.

Inside the bundle sheath cells, the malate is chemically broken down into a smaller organic molecule, and carbon dioxide is released. This carbon dioxide then enters the chloroplast of the bundle sheath cell and is fixed a second timewith the enzyme Rubisco and continues through the C3 pathway.

Advantages of Double-Carbon Fixation

The double-carbon fixation pathway confers a greater photosynthetic efficiency on C4 plants over C3 plants, because the C3 enzyme Rubisco is highly inefficient in the presence of elevated levels of oxygen. In order for the enzyme to operate, carbon dioxide must first attach to the enzyme at a particular location known as the active site.

However, oxygen is also able to attach to this active site and prevent carbon dioxide from attaching, a process known as photorespiration. As a consequence, there is an ongoing competition between these two gases for attachment at the active site of the Rubisco enzyme. Not only does the oxygen outcompete carbon dioxide; when oxygen binds to Rubisco, it also destroys some of the molecules in the Calvin
cycle.

At any given time, the winner of this competition is largely dictated by the relative concentrations of these two gases. When a plant opens its stomata (the pores in its leaves), the air that diffuses in will be at equilibrium with the atmosphere, which is 21 percent oxygen and 0.04 percent carbon dioxide.

During hot, dry weather, excess water vapor diffuses out, and under these conditions plants face certain desiccation if the stomata are left open continuously.When these pores are closed, the concentration of gases will change. As photosynthesis proceeds, carbon dioxide will be consumed and oxygen generated.

When the concentration of carbon dioxide drops below 0.01 percent, oxygen will outcompete carbon dioxide at the active site, and no net photosynthesis occurs. C4 plants, however, are able to prevent photorespiration, because the PEP carboxylase enzyme is not inhibited by oxygen.

Thus, when the stomata are closed, this enzyme continues to fix carbon inside the leaf until it is consumed. Because the bundle sheath is isolated from the leaf’s air spaces, it is not affected by the rising oxygen levels, and the C3 cycle functions without interference. C4 photosynthesis is found in at least nineteen families of flowering plants.

No family is exclusively composed of C4 plants. Because C4 photosynthesis is an adaptation to hot, dry environments, especially climates found in tropical regions, C4 plants are often able to out compete C3 plants in those areas. In more temperate regions, they have less of an advantage and are therefore less common.

CAM Photosynthesis


A second alternative photosynthetic pathway, known as crassulacean acid metabolism (CAM), exists in succulents such as cacti and other desert plants. These plants have the same two carbon-fixing steps as are present in C4 plants, but rather than being spatially separated between the mesophyll and bundle sheath cells, CAM plants have both carbon dioxide-fixing enzymes within the same cell.

These enzymes are active at different times, PEP carboxylase during the day and Rubisco at night. Just as Kranz anatomy is unique to C4 plants, CAM plants are unique in that the stomata are open at night and largely closed during the day.

The biochemical pathway of photosynthesis in CAM plants begins at night. With the stomata open, carbon dioxide diffuses into the leaf and into mesophyll cells, where it is fixed by the C4 enzyme PEP carboxylase. The product is malate, as in C4 photosynthesis, but it is transformed into malic acid (a nonionic form of malate) and is stored in the cell’s vacuoles (cavities within the cytoplasm) until the next day.

Although the malic acid will be used as a carbon dioxide source for the C3 cycle, just as in C4 photosynthesis, it is stored until daylight because the C3 cycle requires light as an energy source. The vacuoles will accumulate malic acid through most of the night.

A few hours before daylight, the vacuole will fill up, and malic acid will begin to accumulate in the cytoplasm outside the vacuole. As it does, the pH of the cytoplasm will become acidic, causing the enzyme to stop functioning for the rest of the night.

When the sun rises the stomata will close, and photosynthesis by the C3 cycle will quickly deplete the atmosphere within the leaf of all carbon dioxide. At this time, the malic acid will be transported out of the vacuole to the cytoplasmof the cell. There it will be broken down, and the carbon dioxide will enter the chloroplast and be used by the C3 cycle; thus, photosynthesis is able to continue with closed stomata.

Crassulacean acid metabolism derives its name from the fact that it involves a daily fluctuation in the level of acid within the plant and that it was first discovered to be common in species within the stonecrop family, Crassulaceae.

The discovery of this photosynthetic pathway dates back to the 1960’s. The observation that succulent plants become very acidic at night, however, dates back to at least the seventeenth century, when it was noted that cactus tastes sour in the morning and bitter in the afternoon.

CAM Plant Ecosystems

There are two distinctly different ecological environments where CAM plants may be found. Most are terrestrial plants typical of deserts or other harsh, dry sites.

In these environments, the pattern of stomatal opening and closing provides an important advantage for surviving arid conditions: When the stomata are open, water is lost; however, the rate of loss decreases as the air temperature decreases. By restricting the time period of stomatal opening to the nighttime, CAM plants are extremely good at conserving water.

The other ecological setting where CAM plants are found is in certain aquatic habitats. When this environment was first discovered, it seemed quite odd, because in these environments conserving water would be of little value to a plant. It was found, however, that there are aspects of the aquatic environment which make CAM photosynthesis advantageous.

In shallow bodies of water, the photosynthetic consumption of carbon dioxide may proceed at a rate in excess of the rate of diffusion of carbon dioxide from the atmosphere into the water, largely because gases diffuse several times more slowly in water than in air.

Consequently, pools of water may be completely without carbon dioxide for large parts of the day. Overnight, carbon dioxide is replenished, and aquatic CAM plants take advantage of this condition to fix the plentiful supply of carbon dioxide available at night and store it as malic acid.

Hence, during the day, when the ambient carbon dioxide concentration is zero, these plants have their own internal supply of carbon dioxide for photosynthesis. Thus, two very different ecological conditions have selected for the identical biochemical pathway.

These two modified photosynthetic pathways adequately describe what happens in most terrestrial plants, although there is much variation. For example, there are species that appear in many respects to have photosynthetic characteristics intermediate to C3 and C4 plants.

Other plants are capable of switching from exclusively C3 photosynthesis to CAM photosynthesis at different times of the year. Photosynthesis by aquatic plants appears to present even more variation. C3-C4 intermediate plants seem to be relatively common compared to the terrestrial flora, and several species have C4 photosynthesis but lack Kranz anatomy.
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