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Yeast Respiration and Fermentation

TEACHER REFERENCE PAGES

GC Instructions

INTRODUCTION Most organisms, including yeasts, use oxygen in a process called cellular respiration. Cellular respiration is the controlled breakdown of carbohydrate to carbon dioxide and water with capture of some of the energy in the form of ATP. The rest of the energy is lost in the form of heat. The first stage of the breakdown is called glycolysis and the second stage is called the Krebs Cycle. During this process, electrons are transferred from the carbohydrates to oxygen in the process called electron transport and water is formed as the final product of electron transport. Electron transport produces a chemosmotic gradient of protons (H+) and positive charges across a membrane and this gradient can drive the formation of ATP. Cellular respiration produces approximately 38 ATP molecules from each molecule of the sugar glucose that is broken down. The carbon that was in the carbohydrate is fully oxidized to form CO during respiration. For glucose, the 6 carbons become 6 CO molecules.
2 2

Table 1. Comparison of respiration and fermentation of glucose in yeast. PROCESS CONDITIONS PRODUCTS FROM GLUCOSE 6 CO + 6 H 2 CO + 2 C H O
2 2 2 2 6

RESPIRATION AEROBIC FERMENTATION ANAEROBIC

AMOUNT OF ATP 38 2

Fermentation, a process that can occur in the absence of oxygen, partially breaks down carbohydrate by glycolysis to capture a small amount of energy in the form of ATP. The initial reactions of fermentation and respiration are the same, but fermentation stops after glycolysis whereas respiration continues into the Krebs Cycle. The carbohydrate leftovers are different depending upon the organism that performs the fermentation; usually one product is more oxidized (electron-poor) than the starting molecule and the other is more reduced (electron-rich). In the case of yeast fermentation, the products from one glucose (C6H12O6) molecule are two molecules of ethanol (C2H5OH) and two molecules of CO2. Human anaerobic (oxygen-free) muscle produces two molecules of lactic acid (C3H6O3). Even though the products are different, each fermentation results in a limited, anaerobic breakdown of carbohydrate with energy release. Since the process does not completely break down the carbohydrate, it does not release much energy that can be captured in the form of ATP. In yeast fermentation, there are 2 ATP molecules produced for each glucose molecule that is fermented. This is a low yield compared to that of respiration, but the ability to perform fermentation allows the yeast to survive and grow in environments where no oxygen exists (see Table 1). Gas Chromatography The major technique that is used to determine the type of organic molecules produced during fermentation is gas chromatography. Gas chromatography (GC) is the separation of compounds in the gas phase, depending on their relative ability to adsorb onto the

column packing and their volatility into the gas phase at the temperature used. The gas chromatograph is a simple, sensitive instrument which can be used to separate and identify about 60% of all known organic compounds. The compounds to be separated are injected into a gas stream which passes through a column at a preset speed. Under a constant set of conditions in terms of temperature, gas flow rate, and column packing and size, repeated injections of a compound elute from (come out of) the column at a nearly constant time from injection. Different compounds elute at different times. One factor which affects elution time is the molecular weight of the compound; heavier compounds move more slowly through the column. Elution time is also affected by polarity and other factors. The column is first injected with known compounds called standards, and their retention times are determined. Then, unknown mixtures of compounds can be injected, and if the known compounds are in the mixture, their peaks can be recognized by their characteristic retention times. A gas chromatograph detects the presence of a compound in its eluate (exiting stream) by means of some property of the compound. One common method used by GC detectors is to compare the conductivity of a heated filament which is placed within a stream of pure reference gas (helium in our lab) to a heated filament placed in a stream of gas containing our sample molecules. When molecules from our sample pass the detector filament, the changes in conductivity caused by temperature changes are converted into electrical signals which appear as peaks on a computer data screen. Peaks seen in the eluate are plotted on a chart, and the integrated peak area is proportional to the concentration of the compound. Many GCs report the integral area of each of the peaks, following the plotted graph of the peaks. An approximate proportionality between peak height and concentration can also be seen (see sample printouts). See Instructions for the Gas Chromatograph. Plant hormones are chemical messengers that are produced in one part of the plant and have a physiological effect on a target tissue that may be distant from the site of production. When hormones reach the target tissue they can: (1) have a direct effect on the target tissue causing a rapid metabolic response; (2) involve the use of a second messenger within target cells; and/or (3) affect transcription of nuclear deoxyribonucleic acid (DNA). Unlike animals, plants have no specialized organs designed solely for hormone synthesis and secretion . Leaves, stem tips, root tips, flowers, seeds, and fruits all produce hormones. Most plant hormones are functional at very low concentrations. Auxins, cytokinins, gibberellins, abscisic acid, and ethylene are the best known plant hormones. All are in some way involved in regulating plant growth and development. Some promote growth by stimulating cell enlargement or division while others inhibit growth by inducing dormancy or promoting senescence. Recently brassinolides, jasmolates, and salicylic acid have been shown to have hormonal function.

Principles of Hormone Function
Often two or more hormones work synergistically. In a classic 1957 experiment, Skoog and Miller provided evidence that auxins and cytokinins work together in the differentiation of plant organs. Using tobacco tissue culture, they showed that when a

tissue culture medium contains low concentrations of auxin and optimal cytokinin levels, then formation of shoots is favored. In contrast, when the culture medium is supplied with optimal concentrations of auxin combined with low concentrations of cytokinins, root formation is favored. Hormones sometimes work antagonistically. Apical dominance is a process in which lateral buds of stems remain dormant as long as the stem apex remains intact. It has been shown that auxin produced in the stem apex is responsible for maintaining lateral bud dormancy by causing cells in the lateral buds to produce another hormone, ethylene, which is a growth inhibitor. During early spring, rapidly growing root tips will generate a high concentration of cytokinin that counteracts the effect of ethylene on the lateral buds of the stem. The lateral buds released from dormancy by cytokinins can then begin growth on their own.

Auxins
Auxins were the first class of plant hormones to be identified. Many auxins, both natural and synthetic, are now known and all have similar effects on plant growth and development. The most widely studied naturally occurring auxin is indol-3-acetic acid (IAA), which is chemically related to the amino acid tryptophan. IAA can be synthesized from tryptophan in intact cells but other synthetic pathways are available. Because auxins can have an effect in very low concentrations, plants regulate synthesis and disassembly of auxin very precisely. Auxins are produced in young shoots and always travel downward in the plant from shoot to root. This polar movement of auxin is not well understood but requires calcium ions (Ca 2 ) and most likely involves special carriers in cell membranes. Naturally occurring auxins promote cell enlargement, are important in tropisms, prevent abscission , promote fruit development, and are involved in apical dominance. Synthetic auxins such as naphthalene acetic acid are used as rooting hormones. Other synthetic auxins include 2,4-D (2,4-dichlorophenoxyacetic acid) and 2,4,5-T (2,4,5-trichlorophenoxyacetic acid) that are used as weed killers. The effect of IAA on cell enlargement has been well studied. IAA stimulates special pumps in the cell membrane of target cells to release H ions into the cell wall, resulting in a pH drop to approximately 5.0 in the cell wall. Enzymes that are pH-dependent then break down important structural bonds between cellulose microfibrils causing an increase in cell wall plasticity . As the cell wall becomes more plastic, water is able to flow in and the cell enlarges. Auxin also may have an effect on transcription of nuclear DNA that can contribute to cell enlargement. Calcium acts as a second messenger in processes involving auxin. Auxin stimulates the release of Ca2 from the vacuole and endoplasmic reticulum in target tissues which affects Ca-dependent enzymes, including kinases , phophatases, and phospholipases.

Other Plant Hormones Cytokinins (for example. gravity. determines sex in cucurbits Jasmonates Involved in response to environmental stresses. unidirectional light will cause auxin to move toward the darkened side of the organ and stimulate enlargement of cells on the darkened side. zeatin. cytokinins work synergistically with auxin in the control of tissue and organ differentiation. isopentenyl adenine) have an effect on cell division. stimulate bolting in biennials. which are growth responses to directional environmental stimuli such as light. This causes the organ to bend toward the light. activate dormant buds Gibberellins Initiate mobilization of storage materials in seeds during germination. causes formation of aerenchyma tissue in submerged stems. As previously mentioned. This effect is often seen in potted plants growing in windowsills. enhance fruit development Cytokinins Affect cell division. control cellular elongation. and touch. control germination of seeds Brassinolides Promote of elongation. Cytokinins are produced in root tips and may be transported in the xylem toward the shoot. delay senescence. involved in apical dominance and various tropisms. cause elongation of stems. . stimulate flowering. stimulates the closing of stomata Ethylene Causes ripening of climacteric fruits. prevention of abscission. stimulate pollen tube growth Abscisic Acid Maintains dormancy in seeds and buds. In phototropism. promotes abscission. stimulate the release of ethylene. can affect tropic curvature Salicylic Acid Activates genes involved with plant's defense mechanisms Auxins are involved in tropisms. promote cell division.Plant Hormones: Roles Hormone Role Auxins Involved in differentiation of vascular tissue.

M ERISTEMS . used.. process. which then lose water. causing the stomata to close. and stimulates formation of aerenchyma (gas transport tissue) in submerged roots and stems.Biology Encyclopedia . organs. Taiz. Evert. 1998. New York: W. Ethylene is the only plant hormone that is a gas. Biology of Plants. Ray F.html#ixzz1ZLC7plEx . and Eduardo Zeiger. and are involved in seed germination. Abscisic acid (ABA). S ENESCENCE George Wittler Bibliography Raven. 6th ed. body. overcome genetic dwarfism. Lincoln. A protein -rich layer just internal to the seed coat. all with a similar chemical makeup. There have been as many as eighty-four gibberellins identified (named GA1 through GA84). function. is probably not involved in leaf or fruit abscission. P LANT D EVELOPMENT . Peter H. has been the most studied. Plant . One role of ABA is the stimulation of stomatal closure. letting chloride ions move out of the guard cells . Brassinolides are plant steroids (many animal hormones are steroids) that may be involved in the light-induced expression of genes. Freeman and Company. Inc. When ABA binds to receptors on guard cell membranes. may determine sex in cucurbits (melon family). 2nd ed. despite its name. MA: Sinauer Associates. the aleurone layer.biologyreference.Gibberellins are a very large class of compounds. and it is involved with leaf abscission. H.cells. During the germination of grass seeds the imbibition (intake) of water stimulates the production of gibberellins by the embryo that diffuse throughout the seed.com/HoLa/Hormones-Plant. and Susan E. The resulting depolarization of the membrane stimulates the movement of potassium ions (K ) ions out of guard cells. Read more: Hormones. responds to gibberellins by synthesizing hydrolytic enzymes that aid in mobilization of stored food in the endosperm for use by the embryo.. is a growth inhibitor that. stimulate bolting in biennials. DNA. Gibberellins promote cell elongation. chloride ion channels open. Eichhorn. but GA3. membrane http://www. animal. called gibberellic acid. Sunderland. Ethylene is also considered a growth inhibitor as it may have a role in causing bud dormancy. causes fruit ripening. SEE ALSO C ELL W ALL . 1999. Plant Physiology.

This tumor was a lump of undifferentiated (having no particular fate) cells. The idea that the cell cycle could be regulated by chemicals was inspired by the relationship between Agrobacterium tumefaciens (a bacterium) and its host plants. the lump could be cured of its infection by either heat shock or by antibiotics and the tumor would continue to grow in a tumor form... This fact indicated that the cells of higher plants are totipotent (capable of becoming meristematic. Once infected. organs. An infection with this bacterium caused a rapidly growing tumor to develop in just about any tissue of suitable host species. and so on.changing its developmental fate). Growing cells in sterile culture was of interest to early cell biologists so that one might study cellular processes without the influences of tissues.Cytokinins The discovery of cytokinins Cell division is a fundamental process of living meristematic tissues. .

It was soon discovered that auxins could initiate the formation of callus in plant tissues too. You observed some callus at the stumps where you applied the auxin, IBA, at 5000 ppm on stems of kidney bean plants. Under good circumstances this can also initiate root formation, demonstrating that dedifferentiated callus tissue can be hormonally induced to differentiate along the line leading to root formation. You also have done a rooting project with Mung beans in which the hypocotyl increased in diameter with callus development, and then nearly burst-open with root development along the vascular traces in the stem. People trying to get tissues to grow in vitro, knew the importance of minerals and vitamins in the medium. They tried various additives to get tissues to grow optimally. Single tissues could grow for at least a limited time, but getting a whole plant to develop from the culture was not possible. One additive that seemed the help a lot was the addition of coconut milk (liquid endosperm). With this one addition and some small amount of auxin, one could regenerate an entire plant from just about any tissue. The search was on to find out what the "magic" of the coconut milk was. Just recently (2001) Folke Skoog died...he and his colleague Carlos Miller tried many possible substitutes for coconut milk. The goal was to find out what chemical could stimulate cell division and be far more reliable that the batch-to-batch inconsistency observed with coconut milk. The nitrogenous base, adenine, had at least some activity in this regard. Fresh herring sperm DNA was totally inactive, but autoclaved herring sperm DNA stimulated the cell cultures. They purified the fractions from this sample until they had a chemical that worked...they named it kinetin...and determined its chemical structure in 1955.

Indeed you can see that this is an adenine derivative (6-furfurylaminopurine). In spite of this, kinetin is not a natural cytokinin. Many nitrogenous bases are modified in various ways in the DNA of organisms, but this particular chemical has never been found to occur without the autoclaving manipulation. This discovery led researchers to start looking for other compounds that would be active in cell division. The concept was based on the idea of structure-activity relationship. Obviously the natural cytokinin must have structure similar to kinetin. Simultaneously the pharmaceutical industry started screening synthetic compounds that would be even more effective than kinetin. The results of those studies are shown above. As you might recall we have been using kinetin and benzyl adenine in various projects in lab. I think you recall from the tissue culture project that we got excellent shoot development in media with some BA and could get excellent callus with a different ratio

of BA:NAA. Indeed the synthetic cytokinin, BA, is used routinely in tissue culture as it is stable to autoclaving and therefore is easy to use in such work. The antagonist discovered has been thought to be a competitive inhibitor for the receptor for cytokinins in plant cells...another structure-activity relationship.

The natural cytokinins
Indeed as the decades passed, the natural cytokinins were found in plant extracts...

Again you will notice how these are adenine derivatives and these can be part of a nucleotide (with added ribose sometimes with the phosphate too). The most common natural cytokinin in plants is trans-zeatin. A graph showing the dose responses of tobacco callus cultures to zeatin and kinetin shows that the callus tissue is far more sensitive to the natural zeatin than it is to the synthetic kinetin...you did this project in class...were your results the same or different?

The cytokinin synthesis pathway The precursor for the side-chains of adenine in cytokinins is generally isoprene. The material made in this case is isopentenyl pyrophosphate: . so the terpenoid biosynthesis pathways are partially shared with gibberellins.

That IS a valid assumption. The pathway continues. the natural cytokinin ribosides can be attached to sugars to form glycosides that have reduced or no cytokinin activity. . The plant also can produce enzymes to cleave the sugar and restore full cytokinin activity. right?! Cytokinins occur in both free and conjugate forms As you can see in the diagram below. ultimately producing IPA and Zeatin. Thus conjugation with sugars and retrieval from these bound forms is a possible pathway in plant cells. I am assuming here that you know how plants produce five-carbon sugars or five-carbon sugar phosphates to do this. more of the enzymes are being found by comparison with bacterial genes.You will notice above that the first steps of the pathway are known only vaguely in plants! But as the genome of Arabidopsis has been searched.

Plants carry out this reaction easily and rapidly. it is also true that the ribosides themselves are a form of conjugation. Cells in culture require the cytokinin to be free..However. but this is an artifact. Cytokinins can be degraded . All studies to date seem to indicate that the free-base has to be cleaved from the ribose too before the compound has any true activity. so ribosides appear to have activity on their own.these sometimes lack the enzymes to cleave the ribose. so supplied ribosides are inactive and the free-base must be supplied in the medium.. The native cytokinins also occur as modified bases in RNA and DNA strands. In fact ciszeatin (a less active form) is found in many tRNA molecules in almost all living cells of all species! The extent to which the free cytokinin pool is altered by conjugation with other nucleotides or released from nucleotide polymers is not clear.

The transported cytokinins can be recovered in xylem sap that exudes from cut stems and this has been found to be in the form of zeatin ribosides. The root-produced cytokinins are transported acropetally to the shoot tip. Indeed this diagram represents the homeostasis of cytokinin pools. .In addition to synthesis and conjugation. The apical bud of plants. the pools of cytokinins can be altered by degradation. Hopefully the discussion above is leading you to thinking about a diagram we have seen before in connection with auxins and gibberellins. Below is how one natural cytokinin is made inactive: Transport of cytokinins is acropetal The primary site of cytokinin synthesis in a plant is most likely the root tip. young primordial leaves and flowers. and developing seeds inside fruits are also known to produce cytokinins.

So what's going on with Agrobacterium tumefaciens? As you recall Agrobacterium infection can cause a cytokinin-induced tumor to develop and the plant can be "cured" of its bacteria by holding it at 42° C. This idea is shown below. .

The genes turn on cytokinin synthesis! The structure of the T-DNA in the Ti plasmid is linearized below. This plasmid is called the Ti (Tumor inducing) plasmid. This piece of prokaryotic DNA has two segments of DNA called the "left border" and the "right border" with genes in between.The Agrobacterium injects a plasmid (naked circular DNA) into the host (in this case tomato) cells... . These "borders" permit recombination of the genes into the host genome.

the gene simply has to be put between the borders of the Ti plasmid (probably along with an antibiotic resistance gene for selection purposes) and let the Agrobacterium inject the gene into the cells for incorporation in the genome. simply demonstrates that the Ti DNA has become a permanent addition to the host genome. . the cells can develop into whole transgenic plants (thanks to totipotency of plant cells!). but the genes remain in the cells. Anytime a scientist wants to insert an engineered gene into a plant cell. This process is briefly outlined below. The fact that the plant can be cured of the bacteria later and the tumor continues to thrive. This realization of course provided the smart researcher with a useful tool for transforming plant cells with foreign DNA.This. so we cut out the cytokinin synthesis gene and replace it with the gene of our research interest. Since the cytokinin over-production genes are absent. Of course we are interested in making a transgenic whole plant. in turn results in the development of a "crown gall" tumor on the plant. The bacteria may be killed.

Here you can see the combinations of hormones that control differentiation.The relative amounts of cytokinins and auxins can regulate the differentiation that can occur from transformed cells. With low cytokinin but lots of auxin you get rooty explants which is what you noticed in our Mung-bean lab exercise. callus proliferates as in the normal Crown Gall form as caused by the normal Ti-plasmid from Agrobacterium. With lots of both cytokinin and auxin. The no hormone control is in the lower left corner. the relative ratio of these two hormones regulates what develops. With low auxin but lots of cytokinin the explants form callus and then produce small shoots. Here you can see an array of cytokinin concentrations and auxin concentrations on callus growth. . A close-up of the four most-interesting of those plates is presented below. Explants just increase in size but maintain leaf morphology in the control. Moving transformed cells among these four hormone concentrations. it is possible to regenerate whole cells from them. So when it is time to regenerate whole plants from engineered cells.

Cuttings produce adventitious roots slowly and require additional auxin to reliably root. Wounding often produces a new branch. Cytokinins delay senescence. The results of applied cytokinins could include release of apical dominance as you demonstrated in kidney beans in laboratory. Apical dominance is released. Those treated with cytokinins .Cytokinins have many roles in plants Cytokinins are known from cytokinin-overproducing mutants to produce additional leaves and branches on the stem. Tumors may form at nodes. The stems and leaves produce additional chlorophyll. Leaf senescence is delayed. You carried out this project at home with isolated wheat primary leaf tips in various solutions of plant hormones. Cytokinins regulate the cell cycle as we learned early in the semester.

. when the tagged leaf is also treated with cytokinin. Of course. remember the auxin dose response. In seedling C.there is such a thing as too much of a good thing! Cytokinins cause nutrient diversion. This is shown in a classic experiment in plant physiology below. . Cytokinin Mode of Action We are only beginning to understand cytokinin modes of action. there is not even the small amount of leakage to the other leaf observed in the control (seedling A)... but here is one idea of where we think this kind of research will lead us.should have demonstrated delayed senescence (stayed green longer). Here you can see in seedling B that the cytokinin-treated leaf on the left attracted the radio-tagged amino acid from the untreated leaf on the right. Cytokinin-treated leaves become "sinks" for nutrients such as amino acids..

this is a self reinforcing cycle. The buds are covered with modified leaves that is the tough scales to protect these tender and delicate structures.which occurs in the axil of a leaf or at the tip of the stem. they influence other effectors that result in cytokinin responses. Terminal Bud Sub Topics • • • • Classification of Buds Terminal Bud Facts about Terminal Buds Significance of Terminal Bud Plant buds are small incompletely developed part of the plant. These two hormones are in constant communication via the plants vascular pathways. The strongly growing root tip produces cytokinen which follows the same pathway back to the terminal bud or shoot where it serves as a strong growth regulator. Strong terminal buds or terminal shoot growth (early in the season) produce a strong auxin signal that does two things. produced by the roots. Unaltered. ultimately ending in phosphorylation of a shuttle protein. When these gene products are. Classification of Buds . This triggers a cascade of phosphorylations of proteins. There it is destroyed. this cycle produces a plant that grows strongly at the branch tips and at the ends of the roots. Hormones and Pathways Hormone changes are no less important but not quite as obvious as food and water balance. A bud remains dormant in unfavorable conditions but is consists of cells which are capable of division when the conditions are favorable. phosporylated. Woody plants typically show strong growth at the branch tip (terminal bud) and the root tip. and auxins. produced by the leaves and buds. It also travels down the pathways to the root tip where it serves as a powerful growth regulator for the root tip. phosphorylates type B ARR proteins. The phosphorylated AHP protein enters the nucleus. AHP. that turn on the synthesis of type A ARR proteins. in turn. There is also a negative feedback loop here to shut down the system when enough phosphorylated ARR is present.Initially the cytokinin signal binds to a receptor's CHASE domain. The two most important for us are cytokinens. It suppresses bud break at all the buds behind it on the branch and stem. As you can see.

In some ornamental plants to make a plant bushier or shorter gardeners removes the apical tip and hence suppresses the effect of hormone. thereby contributing to increase in length of the plant.the cells which posses the capability to divide and grow are known as the meristematic cells. a bud from which leaves (but not flowers) develop are called leaf buds.The terminal buds have this tissue called the apical meristem.The terminal bud produces hormones 'auxin' which inhibits the growth of axillary buds.and is the primary growing point of the stem. while those buds yielding both leaves and flowers in the earliest stages of development are termed mixed buds Terminal Bud Back to Top A bud that developes at the apex of the stem. Plant buds can also be classified according to their internal structures: • Floral buds producing blossoms.Back to Top Plant buds can be classified as: • • • Terminal bud:Buds located at the tip of stem and increases the length of the plant Lateral buds: Buds at sides of the stem.Apical meristems found at the tip of stems and root which contributes to increase in length. In plants.increases the girth of the stem Axillary bud: the buds formed in the angle which leaf forms with the stem and produce branches.which allows the lower dormant lateral buds to develop and permits the side shoots to develop.The terminal bud is dominant as it represses the growth of the lateral/axillary bud below it and this phenomenon is known as Apical Dominance. .

The buds are covered with modified leaves that is the tough scales to protect these tender and delicate structures. .A bud remains dormant in unfavorable conditions but is consists of cells which are capable of division when the conditions are favorable.Facts about Terminal Buds Back to Top Plant buds are small incompletely developed part of the plant.which are formed by the scales covering the terminal bud.The age of the plant can be calculated by counting the scars over the plant.which occurs in the axil of a leaf or at the tip of the stem. Significance of Terminal Bud Back to Top The terminal bud helps in the lengthwise growth of the plant.If we remove the terminal bud the flow of would be redirected to the axillary buds below and force them into growth giving more shoots but delaying flowering.

g.Plant Tissue Culture . tissues. e.These are culture of isolated immature or mature embryos. suspension culture are in vitro cultures of isolated cells and very small cell groups remaining dispersed as they grow in excited liquid media. cell culture is used for in vitro culture of single or relatively small groups of plant cells.The conventional breeding methods are the most widely used for crop improvement.These are cultures of isolated plant organs including cultures derived from root tips..com/Q/What_are_different_types_of_plant_tissue_culture#ixzz1ZLE rQDUw . (d) SUSPENSION CULTURE:. are cultured in vitro to obtain their development as organised structures. plant tissue culture or cell culture:it is the technique of in vitro. But in certain situations.These are culture of tissue arising from disorganized proliferation of cells from segment of plant organs. culture in which isolated plant cells. But in a strict sense. Another term. generally grown on solid medium as amass of cells.answers. embryos. tissue culture denotes the in vitro cultivation of plant cells in an unorganised mass. etc. tissues. Aseptic culture of plant may be of the following types as:(a) ORGAN CULTURE:. suspension cultures. the term tissue culture is applied to both callus and suspension cultures.These are often called cell cultures.stem tips. When organised structures like root tips. tissues as well as organs. In this book. But in general. The term tissue culture is commonly used in a very wide sense to include in vitro culture of plant cells. it is called organ cultures.tissue or callus culture are .leaf primordia or immature part of flowers and immature fruits. (b) EMBRYO CULTURE:. the term tissue culture is used in its broad sense to denote aseptic culture of plant cells. as they represent a lower level of organisation than tissue or callus culture. which is not possible through the conventional methods. shoot tips. and cell culture is often used for callus culture as well. (c) CALLUS OR TISSUE CULTURE:. e..conical flask. organs or even entire plant are subjected to grow in nutrient media in glass containers(tubes. these methods have to be supplemented with plant tissue culture techniques either to increase their efficiency or to be able to achieve the objective.g. callus cultures. and organs.petri dishes) under aseptic conditions. Read more: http://wiki.

and Nutrition on Extension of Lateral Buds in Phaseolus vulgaris L. Kinetin alone showed few significant effects on lateral shoot growth but applied with GA3 it often dramatically increased GA3-induced growth of main stems and laterals. IAA applied alone reduced lateral bud extension slightly. organic sewage. lakes and rivers as well as marine bodies like coastal and deep-water seas. silt. detergents. Deposition of acidic substances causes acidification of water by . D. TIN SHEIN1 and 2. which parallels that found in the literature. New Zealand • Received December 21. The diversity of these results. JACKSON + Author Affiliations 1. agricultural chemicals. on concentration of hormone and on light intensity or nutrition. no meaningful relationships were found and it is concluded that growth of laterals and main stems is dependent on a hormone balance which can be critically modified by a wide range of internal and external factors the nature of which is still to be determined. 1. however. kinetin. both freshwater bodies like ponds. These substances are deposited directly on the water bodies. I. aerosols and other acidic substances released into the atmosphere from the industrial or domestic sources of combustion of fossil fuels eventually come down to the ground. 1972. and indol-3yl-acetic acid (IAA) contained in lanolin were applied in various combinations and concentrations to decapitated stems and petioles and to buds of Phaseolus vulgaris L. when applied with GA3 or GA3 plus kinetin. however. On occasions. GA3 applied alone usually promoted growth of main stems and laterals but this was by no means consistent and occasionally it acted in the opposite way. industrial effluents. it often markedly inhibited the promotion caused by these compounds. The pollution is occurring in all types of water bodies. Lincoln College Canterbury. Abstract Gibberellic acid (GA3). Light. However. these substances also reach the water bodies along with run-off rainwater from the polluted soil. Water pollution and plants Water pollution and plants The water bodies of the earth are being continuously polluted by a variety of sources. Effects of acid deposition Various acid gases. GA3 and IAA acted synergistically to promote and sometimes to inhibit lateral shoot growth. In addition. was shown to be partly dependent on the point of hormone application and age of the plant or bud. but not consistently. Major causes of water pollution are deposition of acid.Interaction between Hormones. oil and heat into the water bodies.

Among green algae. most of the diatoms and green algae disappear below the pH 5. fecal material and other organic material is deposited directly from sewage discharges or is washed along with rainwater into the water bodies.6 while Chlamydomonas acidophila is found in water up to pH of 1.lowering its pH below 6. nitrates and chlorides have been reported to make water bodies like lakes. The roots of macrophytes are generally affected adversely in acidic water and result in poor plant growth. Catharina become the only survivors. In the initial phase of water acidification.0. also become abundant. Eleocharis acicularis. Change in species composition: The number and abundance of acid tolerant species increases while that of sensitive species decreases. Effects of organic matter deposition Large amounts of dead and decaying animal and plant material. Cladophora is highly acid tolerant species and becomes abundant in highly acidic freshwater bodies. This reduces the rate of decomposition of organic matter and. Juncus bulbosus. planktonic green and blue-green algae grow very rapidly causing water blooms. Diatoms and small siliceous phytoplankton populations are highly sensitive to pH changes and species composition of their communities shows highly specific changes with pH change of the water body. Fontanalis. The rapid decomposition of organic matter by these increases nutrient availability in the water. Yellowing of plants is common in polluted water. low pH causes nutrient deficiency and consequent general reduction in abundance of aquatic plants in the affected water body. Euglena and some other unicellular algae are found up to pH of 1.0. Sparganium emersum. Nutrient deficiency in aquatic ecosystem: The decomposing bacteria and fungi decrease in acidified water. the nutrient cycling in the aquatic ecosystem. Eutrophication: Addition of organic matter and its rapid decomposition resulting in increased nutrient supply causes much nutrient enrichment (eutrophication) of water body. The sulphates. filamentous algae grow very fast and form thick mats. Azolla. Potamogeton pectinalis is only aquatic macrophyte found in heavily acdified water. Thus. rivers and ponds acidic in many countries.0.8. therefore. In addition to these many types of hydrophytes like Salvinia.0. Typha latifolia and bryophytes like Polytrichum. Macrophytes are generally absent in extremely acidic water. In such a condition. Decrease of species diversity: Critical pH for most of the aquatic species is 6. Juncus effusus. All this rapid growth of planktonic and free-floating hydrophytes reduces light penetration into deeper layers of water body and . Increase in decomposer microbes: Increased addition of organic matter into the water body results in rapid multiplication and increase in decomposer aerobic and anaerobic bacteria. angiosperm species using dissolved carbon dioxide e. Most important consequnces of such organic matter deposition are as following.g. Eicchhornia etc. The number and variety of aquatic species in the water body generally decreases below this pH. Plants with deep roots and rhizomes are less affected while plants with short root systems are severely affected. However. Anisothecium. At pH lower than 4. Gyceria fluitans.

first into the soil then finally into the water bodies. Cation-active compounds hinder algal growth between 0. Biological oxygen demand (B.B.) of the water. On the other hand.g. pesticides.) of water is defined as the amount of oxygen needed by a unit volume (usually one litre) of water sample to completely decompose the organic matter present in it by microbial activity. it may be more than 400 mg/litre.D. measured at 20oC and tested at least five days after sampling. Binding of membrane lipids and proteins. Denaturation of proteins and thus causing enzyme inhibition in various metabolic processes.0 ppm while non-ionic compounds hinder algal growth between <1. These cause acidification of water. enter the plants through roots or surface absorption. Chemical fertilizers entering the water bodies result in eutrophication by enriching the water with major plant nutrients.O.0 and around 10. pollen germination and growth of pollen tubes. ammonium. cation uptake. The herbicides act directly on aquatic flora but insecticides act indirectly by allowing algal blooms to develop in the water body. Most common effect of these substances is reduced photosynthesis.000 ppm concentrations depending upon the species and the compound. root elongation. Some may uncouple . The effects of these plants on aquatic plants are similar to those of their overdose in foliar application. herbicides etc. Abundant flora after death further increases supply of organic matter in the water body. photosynthesis. Oxygen depletion: Rapid decomposition of organic matter by aerobic bacteria during eutrophication phase consumes much water-dissolved oxygen. The B.submerged flora gradually declines. Most of the detergents that are toxic.1 and 10. These excess chemicals are washed away with rainwater. insecticides. Addition of such detergents into water results in phosphate-enrichment of water. Non-degradable alkyl benzene sulphonates and phosphate-rich detergents interfere with gaseous exchange even in very low concentrations. carbon dioxide fixation. herbicides and insecticides also cause pH changes in the water bodies.O. Common effects of detergents on plants are as follows. gradual decrease of submerged aquatic flora results in reduced oxygenation of water.O. Retardation of plant growth. value of fresh. Destruction of the chlorophylls and cell membranes. Macrophytes are most sensitive to damage by anionic surfactants. Different substances have different patterns of their toxic action. Most of the domestic and industrial detergents contain high (up to 40%) phosphate content. unpolluted water is usually below 1 ppm while in organic matter polluted water. Effects of detergent deposition Various detergents from domestic or industrial use directly released or washed down into the water bodies cause serious effects of plants. Effects of agricultural chemical deposition Many chemical fertilizers. are applied to crops far in excess. decomposition pathways and environmental persistance. Pesticides. Alteration of the absorption maxima of chlorophylls. Both these phenomena together result in increase in biological oxygen demand (B. Many of these fertilizers are acidic in nature e.D.

in general. causes reduced uptake of essential bases. Fly ash forms thick. Fly-ash. The deposition of silt increases the turbidity of water and reduces light-penetration deep into the water. Zn. Lemanea is quite resistant to Zn and Pb. A large amount of these substances deposited on the soil. Ranunculus spp. . Zn-toxicity is less with high Ca for Stigeoclonium and Hormidium while is less with high pH for Hormidium. Species tolerant to turbidity (e. Pb. Efects of silt deposition The top soil removed due to erosion is carried with rainwater or flood water and deposited into the water bodies causing silt deposition in them. Cd-uptake by Nitella and Elodea is less in hard water.g. Myriophyllum spicatum. Change in the floristic composition of the water body is most obvious and direct effect of pollution by such effluents. comes into water bodies indirectly along with surface run-off. While bryophytes appear to be highly resistant to heavy metal toxicity. Silt deposition. For example. Cd. Lemna minor agg. Phytoplankton is particularly affected by silt deposition due to reduction in surface exchange of gases and nutrients.g. S. various organic/liquid effluents and heavy metals e. Photosynthesis and growth in most of the algae is inhibited at 1-2 ppm of Cu++. Se etc.32 ppm. Ceratophyllum demersum.. Impact of metals is reduced in hard.oxidative phosphorylation or inhibit nitrate reductase enzyme. cause specific toxicity effects on the aquatic plants. Sagittaria sagittifolia. are important industrial pollutants of water. Scirpus lacustris) becomes highest followed by species of intermediate tolerance (e. well-buffered freshwater systems. The uptake and bioaccumulation of these substances in aquatic plants is great due to their low solubility in water. Fly-ash increases the alkalinity of water and thus. Reduced oxygen and low temperature also increase metal toxicity. All these phenomena cause death of aquatic plants. Potamogeton perfoliatus. Chelators decrease while methylated forms increase the metal toxicity to aquatic plants. Bioaccumulation of metals is more in mosses than in angiosperms and is usually more in lower plant parts. floating covering over the water surface. There may be synergistic. in all classes of algae. P. Effects of industrial effluent deposition Various inorganic and organic waste products from industries and mining activities are directly deposited into the water bodies. Cholrella is retarded more by Ag than by Cd. Nuphar lutea.g. Callitriche spp. Polygonum amphibium. erectum) while least tolerant species (e. Sparganium emersum. Potamogeton natans. Ni. Organic/liquid effluents disturb the pH of water and depending upon their chemical composition. pectinatus.. Hg or Ni while cell division in the genus is reduced more by Cd than by Cu or Hg at 0.. causing decline in submerged flora. additive or antagonistic interactions between metals regarding their effects on plants. Ti. Cu. Hg. Cholrella is more sensitive to Cu++ than Scendensmus. inhibits growth of aquatic plants.g. This reduces the penetration of light into deeper layers of water body. strains vary in tolerance to metals. Heavy metals usually occur together and with many other types of pollutants so the effects of single metal are usually difficult to interpret. Elodea canadensis.

Reduced oxygenation and high temperature of water causes reduction in the activity of aerobic decomposers. WAter pollution Correlative Inhibition of Lateral Bud Growth in Pisum sativum L. The solubility of oxygen in water is reduced at higher temperature. MANSFIELD + Author Affiliations 1. reduced photosynthesis and general inhibition of enzyme activity with increasing temperature. A. large lakes and particularly near sea coasts causes deposition of oil slicks on water. J. Effects of waste heat deposition Many industries. 1976. aquatic plants show increased respiration. Abstract . particularly thermal power plants. The reduced decomposers result in decreased organic matter decomposition and consequently. Oil may even cause destruction of aquatic flora if it catches fire.: Studies of the Role of Abscisic Acid 1. Oil spills in water are also common during normal transport operations or during accidents involving oil tankers. Department of Biological Sciences. University of Lancaster Bailrigg. Lancaster LA1 4YQ • Received December 15. The aquatic flora and primary productivity of the aquatic ecosystem declines with increasing temperature. The deposition of waste heat into the water body has many consequences for the plants in it. reduced nutrient availability in the water body. T. which have comparatively less primary productivity. Oil depletes oxygen of the water body by consuming dissolved oxygen in oil degradation. and Phaseolus vulgaris L.Mosses) are much reduced. species diversity of the water body declines and heattolerant species gradually become dominant. take water from rivers. Effects of oil deposition Washing of oil tankers and storage containers in many rivers. The heated water is then returned to the water body. C. lakes or sea to cool the heat-producing boilers and equipment. Oil pollution of water body prevents oxygenation of water. In high temperature. Thus. Oil inhibits planktonic growth and photosynthesis in aquatic macrophytes. With increase in temperature. Posted by garg at 6:14 AM Labels: plants. the reduced oxygenation of water adversely affects the aquatic flora. Green algae are mostly replaced by blue-green algae. WHITE and 2.

[4] antioxidant protection through metabolites and enzyme systems to reduce the oxidant load.. glutathione peroxidase. [3] impairment of photosynthesis via changes in Rubisco levels and the guard cells so that the stomata do not track correctly the environment. Riverside. but a high concentration in lanolin (1 mg g−1) did substantially reduce bud outgrowth. Endogenous ABA-like substances in Phaseolus vulgaris. but it was not possible to test this hypothesis by determining endogenous ABA levels in axillary buds because of their small size. most of these changes are likely to have been initiated at the level of gene expression. In the last decade considerable understanding of the biochemical process within plants has been developed. and induced a compensatory growth of the bud on an adjacent node.edu Abstract Plant strategies to survive ozone stress include exclusion or tolerance of ozone. past observations of ozone injury have indicated many physiological and metabolic changes then occur. Many believe that free radicals and other oxidative products. The evidence presented here suggests that ABA is not a correlative inhibitor in Phaseolus vulgaris even though at high concentration it can inhibit the growth of axillary buds. and . such as ascorbate. are responsible for much of the spread of the biochemical alterations. If these processes fail. [2] stress ethylene interactions. CA. evidence suggests the first line of defense is a range of antioxidants. such as hydrogen peroxide. USA. lateral buds on intact plants and lateral buds on plants decapitated 24 h earlier.The possibility has been investigated that abscisic acid (ABA) might act as a correlative inhibitor of lateral bud growth in Pisum sativum and Phaseolus vulgaris. sativum caused appreciable inhibition of their growth. Application of this same quantity of ABA to axillary buds on decapitated plants of Phaseolus vulgaris was without effect. Alterations of the biochemical pathways of plants by the air pollutant ozone: which are the true gauges of injury? Heath RL.l. Once the ozone enters the tissue. University of California. There are obvious chemicals that may account for the changes that are observed. Source Department of Botany and Plant Sciences. were present in similar concentrations in shoot tips. The effects of applied ABA suggested that it might be involved in the mechanism of correlative inhibition in Pisum sativum. Currently there are several hypotheses regarding a response of plants to ozone fumigation: [1] membrane dysfunction and alteration of purpose. heath@ucr. formed in plant leaves under ozone exposure. suggesting signal transduction. detected by bioassay and electron capture g. Application of ABA in small quantities (2μg) to axillary buds on decapitated plants of P. and [5] general impairment or disruption of metabolic pathways. superoxide dismutase.c.

commercial applicators. The dramatic strides in understanding the genetic make-up of plants. What is now known about how varied biochemicals and their pathways are changed upon ozone exposure will be discussed. . Arthur Lamey Extension Plant Pathologists The interactions between plants and disease organisms are complex. examples of common North Dakota diseases were selected from both the commercial farm and from the home garden. home gardeners and others who want more information on how plant diseases develop and are managed. subsequent events occur which are highly suggestive of systemic acquired resistance. Plant Diseases Development and Management EB-31 (Revised). This illustrates the first important concept: there are two basic types of diseases. but more slowly than ethylene. The primary set of metabolic reactions that ozone triggers is thought to be "wounding" responses with a secondary response of senescence. as can many fungi and other infectious living organisms. To serve these diverse groups. HOW DISEASE DEVELOPS WHAT IS A DISEASE? A disease is any abnormal condition that damages a plant and reduces its productivity or usefulness to man. other defensive indicators. Various circulars available at offices of the NDSU Extension Service provide detailed information on many of these diseases. Furthermore. McMullen and H. February 2001 Marcia P. If overwhelmed. and signal transduction/control over the last few years will only accelerate in the future. and spread their signaling effects more widely in the plant. air pollution can cause disease.catalase. gene control. Confusion can be reduced by learning a few basic concepts and principles of how diseases develop and how they are managed. non-infectious (abiotic) and infectious (biotic). and commercial growers and home gardeners alike may have difficulty understanding plant diseases. Under this definition. such as salicylic acid and jasmonic acid. This publication presents these concepts and is dedicated to commercial growers. We need now to have an understanding of those events that can be translated into more detailed schemes of how ozone alters much of the basic metabolism of plants and how plants counteract or cope with ozone. tend to increase.

these leaves are yellow with green veins. Bright . some tree canker organisms commonly infect trees stressed by drought or extreme cold. Iron chlorosis also is common on certain soybean varieties. Nutrition Nutrition is a frequent cause of non-infectious disease. parasitic organism (an organism that gets its food by attacking other organisms). The effects of stem rust and root rot on small grains are greater when plants are moisture stressed (deficient in water). are common. Moisture Deficient or excessive moisture (water) can cause disease. Excess moisture also has adverse effects. In addition. leaves may turn brown and become brittle as well. caused by iron deficiency. this stress may predispose (weaken) plants to infection by infectious organisms or increase the effects of infectious disease. In oats. Lime-induced chlorosis is common in our alkaline soils because the iron in the soil is not readily available to plants. plants that are deficient in nitrogen develop a general yellowing. Either too much (excess) or too little (deficiency) can cause problems. Iron chlorosis occurs in many North Dakota trees and shrubs. beginning with the lower leaves and progressing upward. but these are rare in North Dakota. causing yellow leaves and stunted growth. Zinc deficiency is common on dry beans and fairly common on flax.TYPES OF DISEASES NON-INFECTIOUS (Abiotic) Non-infectious diseases are caused by some environmental factor that produces an abnormal plant (Table 1). stressed or wilted plants. Excess trace elements may also cause growth problems. oak. Iron chlorosis is recognized by progressively smaller leaves on the new growth." Other Meteorological Conditions High soil temperatures early in the season may injure or kill plant tissues at the soil surface. such as suffocation of roots due to lack of oxygen or predisposing plants to water mold infections. Trace element deficiencies such as iron chlorosis. Moisture deficiency produces stunted. one that has an abnormal appearance. especially silver maple. this is referred to as "blast. For example. For example. this is called heat canker. but are abiotic in nature. Noninfectious diseases are not caused by a living. that is. For example. and spirea. Temperature Frost is a common problem in spring and fall. affecting tender farm crops and garden vegetables. When iron chlorosis is severe. Extremely high temperatures in summer can also cause problems. resulting in a constricted stem. heat sterility in small grains is common in North Dakota.

Many live on dead or decaying organic matter and are called saprophytes. and other types of tissue death. Most pathogenic fungi produce spores which serve to reproduce and disseminate them. excessive growth. and canola. not all fungi are parasites. root rots of small grains. The sooty molds seen on . Spores function similarly to the seeds of higher plants. Bronzing of beans caused by ozone is common in the state. Symptoms are also useful in identification of a disease. stunting. Other spores develop in specialized fruiting structures. Symptoms are visible abnormalities such as wilts. When lightning strikes the ground it may kill plants in somewhat circular patches up to 50 feet in diameter. cereal smuts and head scab of small grains. anthracnose of muskmelon. These structures are called signs of the pathogen and are useful in field identification of disease. rots. The pathogen can spread from a diseased plant to a healthy plant. white mold or Sclerotinia of dry beans. Mushrooms that spring up in lawns are among the most spectacular saprophytic fungi. The source of the ozone is not known. There are also many inconspicuous ones that rot organic matter. Fungi Fungi are the most common pathogens in North Dakota. Toxic Chemicals Toxic chemicals injure plants. peony blight. The plant attacked is called the host plant. sunflower and dry beans. There are five common groups of pathogens (Table 1). resulting in sunscald. INFECTIOUS (Biotic) Infectious diseases are caused by organisms that attack plants and get their nutrition from them. Salt may damage or kill farm crops growing in saline seeps. Examples of common pathogenic fungal diseases in North Dakota include: rusts of small grains. Some spores are formed in masses.sun. organisms that get their food from other living organisms. Cercospora leafspot of sugarbeet. apple scab. The organism causing the disease is called a pathogen. like the orange pustules of rust fungi. They produce tiny thread-like filaments called hyphae. sugarbeets and dry beans. Septoria leafspot of tomato. early blight and late blight of potatoes. sunflowers. Air pollution also damages vegetation. road salt may severely damage boulevard trees and other vegetation. A few other kinds of micro-organisms may cause plant disease but are not common in North Dakota. powdery mildews of ornamentals. However. high temperatures. and plum pockets. Most fungi that cause plant diseases are parasites. and strong dry winds may suddenly desiccate (dry) leaves of crops and garden plants. or abnormal color. tree cankers.

carrots develop yellow tops and hairy roots. Phytoplasmas cause growth abnormalities such as witches' brooms (a broom-like mass of plant branches) or excessive tillering (stooling of small grains). cucumber mosaic. have no defined shape and can only be seen with an electron microscope. The pine wilt nematode is unusual in that it invades the vascular (water-conducting) tissues of pine trees instead of attacking the roots. bean common mosaic. but a few can be seen with the naked eye. others are transmitted (carried) by insects and by eriophyid mites.wheat heads at harvest also are saprophytic. Examples of virus diseases that can cause serious losses are wheat streak mosaic. and have a protein coat covering the core. They are usually systemic in the host (distributed internally throughout the host) and are transmitted by leafhoppers. The disease produces greenish flowers on flax. Nematodes Nematodes are tiny roundworms. They are the soybean cyst nematode and the pine wilt nematode. fireblight of apples and related plants. They can be seen only with a microscope. bacterial blights and black chaff of wheat and barley. Aster yellows produces witches' brooms and greenish flowers on marigold. Purple top of potato is often accompanied by the formation of small aerial tubers in the leaf axils (the point where the leaf joins the stem). potato virus diseases. Viruses are usually in the form of rods or spheres and alter the activities of the host to manufacture more virus. Two potentially serious nematode diseases occur in neighboring states but are not yet confirmed (2000) in North Dakota. and bladder-like pods on canola. Some parasitic forms attack plant roots and can cause severe damage. Most are saprophytes. Most can be seen only with a microscope.000 times smaller than the tiniest living cell. the basic unit of heredity. barley yellow dwarf. Our most common phytoplasma is aster yellows. Bacteria Bacteria are tiny one-celled organisms that multiply by cell division. Examples of common bacterial diseases include bacterial blights of dry beans. living on the already ripe or senescing glumes and awns. ring rot and blackleg of potato. Nematode problems are common in warm climates but are rare in North Dakota. "purple top" on potato and tomato. Most viruses have a core of nucleic acid. . angular leafspot of cucumber. and squash mosaic. tobacco mosaic. Some viruses are transmitted mechanically (by contact with another plant. bacterial wilt of cucumber and muskmelon. Viruses Viruses are 1. but there are a few common and serious bacterial pathogens that attack North Dakota plants. and bacterial speck and spot of tomato. Reproduction is by formation of eggs. Phytoplasmas Phytoplasmas lack a rigid cell wall. or contaminated workers' hands or tools).

The disease triangle — disease develops only when all three factors are favorable. Flax rust develops only when a suitable combination of rust race (pathogen) and variety (host) interact under environmental conditions favoring this disease. the flax rust pathogn attacks only flax as a host. Many diseases are favored by humid or rainy weather and may be more common and severe in years favoring good crop production. and different races of flax rust attack different varieties of flax. (4KB illustration) INFECTION BY PATHOGENS Pathogens can infect plants in several ways (Figure 2). WHAT DOES IT TAKE TO PRODUCE DISEASE? Three factors interact to produce disease. . For example. and how they survive between crops.DISEASE DEVELOPMENT This section discusses how plant pathogens produce disease: how they infect the host. The environment often limits disease in North Dakota and is the reason that some diseases rarely occur and others occur sporadically. the host. Viruses. fungi. the pathogen. Figure 1. how they are disseminated (spread). and bacteria also enter through wounds. and the environment (Figure 1). disease will not develop. If any one of these three factors is unfavorable or missing. Many fungi can penetrate the leaf surface directly without wounds or stomata. Fungi and bacteria may penetrate (enter) through natural openings such as the stomata — these are tiny "breathing" pores in the leaf that allow the exchange of gases (Figure 3). Bacteria frequently enter through hail wounds. how they reproduce.

Wind Wind disseminates fungus spores from plant to plant in a field or across fields. see Table 1. and establish infection. . especially foliar (leaf) pathogens. animals and birds. Many pathogens. The aster leafhopper. Tobacco mosaic virus and potato virus X are common examples.) (30KB illustration) Some viruses and most phytoplasmas are carried by insects. Leaf rust is windborne from the major winter wheat areas of Kansas. (Shown in cross section — edge of cut open leaf in foreground. wheat stem rust. Nebraska and Oklahoma (Occasionally it survives the winter in North or South Dakota). water. penetrate the host. Some typical pathogens are illustrated. Some viruses are transmitted mechanically by the rubbing together of leaves or by humans touching diseased and then healthy leaves. Pathogens such as the wheat leaf rust. (14KB illustration) Figure 3. and barley stem rust pathogens are spread long distances by the wind (Figure 4). Insects that carry and transmit plant disease organisms by their feeding are called vectors. This is why wet or humid weather is so important in the development of many fungal and bacterial diseases. especially sucking insects such as aphids and leafhoppers. Stem rust is windborne from Mexico and Southern Plains states to wheat and barley crops in the Northern Great Plains. but they are not shown on the same size scale. For relative sizes. need a film of water on the plant to begin growth. man. insects. for example. is the vector of both the aster yellows phytoplasma and the oat blue dwarf virus. Methods of infection by pathogens. Many fruit tree viruses are transmitted by grafting. DISSEMINATION OF PATHOGENS Pathogens are disseminated (spread) by wind.Figure 2. Diagram illustrating some of the ways pathogens can infect a leaf.

Man can locally disseminate bacterial blights. the soybean cyst nematode can be disseminated in the feces of birds. Verticillium. Insects also spread these pathogens from plant to plant. Animals and Birds Animals and birds also may disseminate pathogens. and luggage for prohibited pests at ports of entry. Man Man spreads pathogens and weeds over long distances.Figure 4. could be introduced on contaminated farm implements from infested areas in neighboring states. (8KB illustration) Insects Insects are important in carrying viruses and phytoplasmas from southern areas. For example. The soybean cyst nematode. animals may spread water-borne pathogens by walking through an infected crop when the plants are wet. only the striped cucumber beetle is common in North Dakota. International plant quarantines attempt to prevent this dissemination by authorizing inspection of planes. The gardener who brings fruit in his or her luggage and the commercial grower who brings seed of a high performance crop in his pocket may introduce a new pathogen or other pest. The Mediterranean fruit fly as well as many weeds and pathogens are frequently intercepted at ports of entry. Bacterial wilt of cucumber and muskmelon is disseminated by both the striped and spotted cucumber beetles. Septoria leafspot of tomato and the bacterial blights of dry beans are common examples. rust and anthracnose of dry beans by cultivating the crop when it is wet. Rain and splashing water can disseminate many fungi and bacteria. not yet reported (2000) in North Dakota. Water Water can carry pathogens from field to field. Long distance spread of wheat stem rust and wheat leaf rust spores from their overwintering areas. much of its local spread and buildup has been on elm firewood collected from diseased trees. ships. cars. Dutch elm disease was originally introduced into the United States on elm logs. . Water flowing over the surface of fields spreads disease organisms such as Sclerotinia (white mold). Common insect vectors include aphids for the barley yellow dwarf virus and the aster leafhopper for the aster yellows phytoplasma. Rain and wind form numerous tiny airborne water droplets called aerosols. and downy mildew. Man can also disseminate pathogens and other pests locally. Many bacteria are disseminated long distances in wind-driven aerosols.

These include the leafspot pathogens of wheat and barley which survive on stubble. Mild Climates The cereal rust pathogens usually do not overwinter in North Dakota but survive year-round in the southern United States and Mexico and are carried north each year by wind. For example. Soil Many pathogens form resistant structures that survive long periods of time in the soil. Insects and Mites The bacterial wilt pathogen of cucumber and muskmelon is suspected to overwinter in the digestive tract of cucumber beetles. When this is over the winter. Many potato pathogens are carried on or within the tubers. it is called overwintering. roots and corms provide a mode of survival for many pathogens.SURVIVAL OF PATHOGENS In North Dakota we are primarily concerned about pathogen survival between crop seasons. and the tomato Septoria leafspot pathogen which survives on the old dead vines. Seed and Vegetative Plant Parts The smuts of small grains survive on or in the seed and survive in storage as long as the seeds remain viable (can germinate). Destroying or burying this crop refuse reduces next year's disease potential. tomatoes and other crops survives for at least several years in the soil. Vegetative plant parts such as tubers. HOW DISEASES ARE MANAGED . and the sunflower downy mildew pathogen survives for at least 14 years as resistant spores in the soil. The Verticillium wilt pathogen of potatoes. The white mold pathogen of dry beans and sunflower survives for six to eight years or more as resistant bodies in the soil. the Aphanomyces root rot pathogen of sugarbeets survives for over 20 years as resistant spores in the soil. Plant Parts Pathogens may survive on crop refuse. The wheat streak mosaic virus overinters in wheat curl mites that survive on winter wheat and some perennial grasses.

management practices prevent or delay the introduction of pathogens or reduce initial pathogen populations and retard their subsequent increase. State and local quarantines are used to keep black wart of potato and the golden . Protectant fungicides and resistant varieties slow down the development of pathogen populations. Non-infectious diseases are not discussed as their management involves a remedy of the physical factors that induce them. in the case of dry bean diseases three different types of fungicides are used for management of each of three major diseases: rust. the type and the timing are important. For head scab of wheat and barley. and white mold. and protection (including the use of fungicides). Planes. trucks and luggage are checked at ports of entry to prevent the introduction of pathogens and other pests into areas where they do not occur. eradication. EXCLUSION Exclusion means exclusion of pathogens. QUARANTINE International quarantines are familiar to anyone who has traveled overseas. This is essential to distinguish infectious from similar appearing non-infectious diseases as well as to correctly identify the pathogen involved in infectious disease. Cultural practices also can reduce the pathogen's population. host resistance. For example. cars. In short. bacterial blights. This information is needed to develop a management program that attacks the pathogen at the weakest point in its life cycle. Correct identification of the pathogen is essential to know the pathogen's life cycle and how it relates to the cycle of disease development. In each case timing is important and application must be started before the disease is widespread. The four basic methods of infectious disease management are: exclusion. at early heading for barley and early flowering for wheat. Pathogens can be excluded (or kept away) from hosts by quarantines that prevent their introduction. These four methods reduce pathogen populations or slow their development. and by use of seed stocks certified to be pathogen-free or within certain prescribed tolerances for low levels of pathogens. Sound management is based on correct diagnosis.This section focuses on management of infectious diseases. When fungicides are used. timing of fungicide application is critical as well. Many disease organisms would flourish in our state if they were introduced.

Fruit trees and potatoes are commonly indexed for viruses. For example. Only those trees free of virus are used as sources of budwood for graft propagation. Indicator Hosts Indicator hosts are plants that produce rapid and distinctive symptoms when inoculated with a virus. INDEXING Indexing involves laboratory or greenhouse tests to determine infection by pathogens in vegetatively propagated plants such as potatoes and fruit trees. They are used to detect specific viruses in individual plants. carnations and potatoes. Crops are field inspected and must meet certain tolerances to be certified. International quarantine has prevented new introductions of pathogens from other countries. Plant parts found free of pathogens are used for further vegetative increase and propagation.nematode restricted to a few localized areas of the eastern United States. This technique is used on chrysanthemums. Certification may be done in conjunction with indexing. Sometimes the seed crop is grown in an isolated area to reduce disease potential. a western county isolated from the rest of the potato production in North Dakota. It is also used to . Indexed nursery stock is commonly used for orchard plantings. These antibodies are produced in the blood of rabbits (usually) that have been immunized with a specific pathogen. Only the healthy materials are saved for further increase. foundation seed potatoes are grown in Golden Valley County. Many seed potatoes are now produced in greenhouses. as this is the only practical way to control fruit tree virus disease. Serology Serology is the use of specific antibodies present in the antiserum of warmblooded mammals. The crop is grown from seed produced under carefully controlled conditions. SEED CERTIFICATION Seed certification is used to certify that potato seed tubers and seed of dry beans have low levels of pathogens or are pathogen-free in the case of potato ring rot. This procedure is used to index barley and potato seed stocks for specific viruses. Culturing Plant parts to be used for increase are laboratory cultured to determine if they are infected with pathogenic fungi or bacteria.

use of resistant host varieties is the practical solution. The embryo test is not available for use on wheat seed. However. Crop rotation helps keep populations of these pathogens at low levels. the percentage of infection. If weather favors disease development. Many highly specific and sophisticated tests are available now. There is one precaution. If the loose smut infection exceeds 1 or 2 percent the seed lot should be treated with an effective systemic fungicide prior to planting. this term may be used when the pathogen is not completely eliminated but the populations are greatly reduced. and Septoria leafspot of tomato attack only one host. The sunflower downy mildew pathogen. so it is difficult to eliminate by rotation. the sugarbeet Aphanomyces pathogen and the white mold organism survive many years in the soil. Cercospora leafspot of sugarbeet. and populations of the pathogen increase when the same host is grown repeatedly on the same land. Crop rotation is an effective tool for reducing many pathogen populations. dry bean rust. however: the grower must consider nearby areas as well. ERADICATION Eradication means elimination of the pathogen. Embryo Test For Barley Loose Smut Detection The embryo test determines if loose smut is present and. kidney and black beans show partial resistance to white mold. only a few hybrids are resistant to all races of the mildew fungus. In the case of sunflower downy mildew. Disease organisms may spread from nearby fields or garden plots if disease was present in those areas the previous year. The pathogen that causes Verticillium wilt of tomato survives several years in the soil and also attacks many other garden vegetables. some navy.detect some bacteria. Pathogens such as the fungi that cause tan spot of wheat. some pathogens survive many years in the soil and are not affected much by normal crop rotations. In actual practice. In the case of Verticillium wilt in tomato. Long rotations may be necessary but often are impractical. the entire field or garden plot may eventually become diseased. . CROP ROTATION Crop rotation involves growing different crops in the same field or plot in succeeding years. if it is. Disease can be expected to develop first in the area next to last year's crop. In the case of white mold.

. It was hoped that eradication of barberry from the Upper Midwest in the 1930s would break the pathogen cycle and eliminate the stem rust fungus. burying or sending to the landfill. To manage apple rust. Burial by tillage reduces the inoculum of the wheat tan spot pathogen. and 2) the sexual phase of the fungus was eliminated. without requiring the barberry for the overwintering stage. Examples of alternate hosts include common barberry (not ornamental barberry) for wheat and barley stem rust. is essential to overwintering of many rusts in northern climates. which slowed down the development of new rust races. the barley spot blotch pathogen." or PP-469 Rev. These spores are wind-blown thousands of miles north to the Upper Midwest every year (Figure 4). and juniper for apple rust. All diseased tomato vines and refuse should be removed. "Plant Disease Control in the Home Garden. who must use fungicides or resistant varieties for management of apple rust when weather is wet in spring and early summer. Diseased leaves and vines also can be composted if the compost is allowed to heat sufficiently. After most barberry had been eradicated. Tillage is sometimes used to bury the refuse. The second host. This is not feasible for the homeowner. buckthorn for oat crown rust. Some of these same rust fungi reproduce indefinitely without the alternate host in warm climates. the barberry eradication program had two very important accomplishments: 1) stem rust infections started later in the season. commercial growers try to remove all junipers within two miles of their orchards. Elimination of buckthorn near oat fields produces similar results. Nevertheless.ERADICATE ALTERNATE HOSTS Many rust fungi require two hosts to complete their life cycle. The rust fungi's sexual phase occurs on these alternate hosts. It was determined that stem rust survives year-round in Mexio and the Gulf Coast in the summer spore stage. and apple leaves that had apple scab should be raked up and detroyed by burning. SANITATION Sanitation is the removal of crop refuse.. The quantity of a pathogen available to produce infection is called the inoculum. stem rust still occurred. as described in Extension Circular PP-737 Rev. and many garden pathogens. "Home Garden Disease Control Begins This Fall. called the alternate host." .

Seed . or development is greatly slowed down. HOST RESISTANCE Resistance is the ability of a host to resist infection by a pathogen. Many foliar pathogens are extremely variable and produce billions of spores that are disseminated great distances by the wind. Consequently. There are two types of host plant resistance: race specific resistance and general resistance. This results in disease outbreaks and a continued need for plant breeding programs. However. General resistance usually does not exhibit as high a level of resistance as race specific resistance. Homeowners who have the Verticillium wilt pathogen in their garden soil must grow a resistant tomato variety to manage the disease. Storing potatoes and other vegetables in cold storage protects against infection because it is too cold for many pathogens to develop. General resistance is usually a stable type of resistance that is effective against all races of the pathogen. but new races may not become widespread as quickly. PROTECTION Protection means protecting plants from infection. Soil-borne pathogens (root and vascular wilt pathogens such as Verticillium) are also variable. Resistant varieties are favored by commercial growers and gardeners when they are available. Resistance has been the best and most cost-effective method of managing stem rust and leaf rust of wheat.When Septoria leafspot of tomato is severe. Some foliar (leaf) pathogens may rapidly develop new races quite capable of attacking certain types of host resistance. Race specific resistance usually provides a high level of resistance. new races of some foliar pathogens may become widespread in a short period of time. picking off badly diseased leaves (sanitation) before spraying with a fungicide helps reduce the inoculum and improves fungicidal control. general resistance slows down disease development compared to that on a susceptible variety. but it fails when new races of the pathogen develop. In the late 1990s new races of wheat leaf rust developed that attacked some of the previously resistant wheat cultivars.

A prime example is the cucumber beetle. MANAGING INSECT VECTORS Many insects carry disease organisms such as viruses and bacteria. Powdery mildew is a common problem on alpine current. Mature tubers are less prone to infection. This includes hilling the soil around the plants to reduce the chances of late blight spores coming into contact with the tubers. CULTURAL PRACTICES Time of planting may help plants escape infection. Pruning shrubs and trees to allow better air circulation and sunlight penetration may help reduce powdery mildew in shady locations. The late blight fungus on the tops will be minimal at harvest and tubers will be mature. Winter wheat is planted in September after the destruction of volunteer wheat. The wheat curl mite is the vector of the wheat streak mosaic virus. The mulch produces a more uniform soil moisture. a noninfectious disease that develops under conditions of drought or fluctuating soil moistures.potatoes are grown in isolated areas where aphid populations are low (exclusion) and thus easily managed (protection). A plastic mulch used on tomatoes greatly reduces blossom end rot. lilac. Most powdery mildews are favored by high humidity. Destruction of volunteers prior to winter wheat planting destroys the green bridge that wheat curl mites survive on between the summer and fall crops. Managing these insect vectors may reduce the chance of disease. The vines should be killed several weeks before harvest by using approved vine killers or chopping the vines off at ground level. HANDLING PRACTICES Development of potato late blight in storage can be minimized by proper handling practices during the growing season. so beans should not be cultivated when they are wet. Delayed planting of winter wheat also reduces the risk of survival and buildup of the mite in the fall of the year and exposure of the wheat crop to high wheat curl mite populations. The bacterium is carried from plant to plant by the beetle and . vector of the bacterial wilt pathogen of cucumber and muskmelon. Many dry bean and garden bean pathogens are disseminated in water. this minimizes aphid-borne virus infection. roses and shaded lawns.

peppers and many other commercial and garden crops. The dicarboximide fungicides iprodione (Rovral) and vinclozolin (Ronilan) also are protectant fungicides that provide good control of Sclerotinia (white mold) on certain field crops. chlorothalonil (Bravo). Some new fungicides have limited therapeutic (curative) properties." Protectant Fungicides Protectant fungicides work on the plant surface to prevent spore germination or kill developing fungus hyphae before the host plant is penetrated and infection becomes established. Protectant fungicides commonly used on North Dakota farms include maneb (Maneb 80. chlorothalonil (Daconil. and copper fungicides. Most protectant fungicides cannot stop development of a pathogen once infection occurs. and copper fungicides. maneb. and seven to 14 days for many other pathogens. Weed management is essential around plantings of potatoes. Ortho Multi Purpose Fungicide). good aphid management is essential for raising virus-free seed potatoes. melons. Then frustrated growers . "Field Crop Fungicide Guide. Current fungicide recommendations are given in Extension Circular PP-622. potato late blight and sugarbeet Cercospora leafspot. If a protectant fungicide is applied after infection has occurred but before leafspots show. mancozeb (Dithane DF." and Circular F-1192. "Insect and Disease Management Guide for Woody Plants in North Dakota. and systemic fungicides are taken up by the plant tissues and then function to prevent infections. weeds and other hosts that can serve as a reservoir for both vectors and pathogens must be managed. cucumbers. A good program for managing the cucumber beetle.overwinters in the beetle. thiram. Agri Tin). the disease is not cured but continues developing and symptoms appear. started as soon as the plants emerge. In addition to successful management of insect vectors. Manzate 200 DF. Leafspots become visible a number of days after infection: four to five days for tan spot of wheat. Protectant fungicides commonly used on North Dakota home gardens or ornamentals include captan (Orthocide). FUNGICIDES Protectant fungicides act on the plant surface to protect against infection. Similarly. Penncozeb DF). Maneb 75 DF). triphenyl tin hydroxide (Super Tin. tomatoes. will prevent serious losses from the bacterial wilt disease. mancozeb.

and if those crops have good yield potential. but uniform coverage is still essential. Note that spraying after infection even though symptoms are not yet evident. if environmental conditions are favorable. If necessary. Only two or three tip leaves were uninfected at the first spraying. Protectant fungicides are redistributed on the plant by dew and rain. These leaves were all that was protected against infection and are all that will be left after disease has defoliated the rest of the plant. If the gardener had sprayed when the first few spots showed on the lower leaves. The disease continues to develop on the middle leaves resulting in defoliation (loss of leaves). Since some upper leaves were already infected before spraying started. Gardeners sometimes begin spraying their tomato plants with a protectant fungicide such as Ortho Multi-Purpose Fungicide when all the lower leaves on their tomato plants are dead and the middle leaves already are heavily spotted with Septoria leaf spot. The application must be made early enough so that the spray droplets have dried before the rain begins. Applying too late will mean that rust infections will already have occurred and new spore formation will not be stopped. Rust diseases have a tremendous capacity to reproduce in a short period of time. Waiting too long usually results in failure. some lower leaves might have been lost to Septoria. These compounds help to wet the leaf and to bind the fungicide to the leaf. will not prevent disease development. growers should apply protectant fungicides to the crop prior to rust formation on the healthy leaves. Effect of spraying a protectant fungicide before or after an infection period (disease-favoring weather). (15KB illustration) . Rapid development of these diseases is favored by very susceptible varieties and rainy. A fungicide should be applied before a rainy period to provide protection during the rainy period. Protectant fungicides must be applied so that thorough coverage of the foliage is obtained." Protectant fungicides do not cure infections and will not work unless the application is timely (early enough). Timely application of protectant fungicides also is critical for crop diseases. such as rust of dry beans or leaf rust of winter wheat. Protectant fungicides are often combined with a spreader-sticker which reduces wash-off of the fungicide during rainfall. humid weather. another application can be made after the rains are over (Figure 5). Spraying before an infection period protects against infection. A few examples will illustrate this. Figure 5.and gardeners may claim that the fungicide "didn't work. but the rest of the plant would have remained healthy. If rust is detected on these susceptible crops in a given area. too. symptoms may develop on these leaves.

Phenylamide (acylalanine) fungicides Metalaxyl (Allegiance) and mefenoxam (Ridomil Gold. Seed treatment is discussed in extension Circular PP-447. This therapeutic effect of sulfur is an exception to the general rule that protectant fungicides do not cure infections." Seed treatments are used to control both seed-borne diseases and soil-borne diseases that can cause death of seedlings. the sterol inhibitors. thiabendazole (Mertect. they are effective only if there is complete coverage of the entire plant. They function as locally systemic fungicides and usually do not move into new foliage. Phytophthora (potato late blight pathogen) and the downy mildew organisms. Sulfur is a protectant fungicide that effectively reduces powdery mildew on a number of crops. including sugarbeets. The white mold fungus infects dry beans by means of airborne spores which must begin growth on dead plant tissue. and some have a therapeutic effect. Benzimidazole fungicides Benomyl (Benlate) and thiophanate methyl (Topsin M) are currently registered for control of white mold (Sclerotinia) on dry beans and give good control if properly applied at the right time. Since these fungicides cannot move down the plant. They include some seed treatment fungicides such as carboxin (Vitavax). "Seed Treatment for Disease Control. they are not washed off by rain or decomposed by sunlight. Systemic fungicides Systemic fungicides are taken up (absorbed) by the plant. Most sterol inhibitors and strobilurins are locally systemic. Arbotect). The site of this dead tissue is the dead blossoms or dead lower leaves. These fungicides move upward in the plant but cannot move down the leaf or stem. Canopy penetration is essential to good control. The therapeutic effect ranges from 24-36 hours for the benzimidazoles and up to four days for some sterol inhibitors. garden and dry peas and ornamentals. the phenylamides and the strobilurins. the benzimidazole fungicides. . Benzimidazoles are commonly used as foliar fungicides and include benomyl (Benlate). Locally systemic fungicides have several advantages: they protect both sides of the leaf even if only one side was sprayed.Protectant fungicides are used both as foliar fungicides and as seed treatments. Apron XL) are systemic fungicides with activity against water molds and related fungi such as Pythium. the interval between sprays may be longer than that with the protectant fungicides. Sulfur also has some eradicant action against established powdery mildew infections. and thiophanate methyl (Topsin M). Oxadixyl (Anchor) is a systemic seed treatment fungicide with similar mode of action to metalaxyl and mefenoxam. and then growth spreads to green tissue. TBZ.

something that is extremely rare with most classes of fungicides. and they are used at very low dosages. a phenylpyrrole class of chemistry that is an environmentally friendly product registered for seed treatment on many crops and also as a potato seed piece treatment. which is a different mode of action from that of other fungicides. Some strobilurins such as azoxystrobin are locally systemic. many for up to four days after infection has initiated. for home garden use). Another new chemistry is fludioxanil (Maxim)." Some are locally systemic and many have an extremely broad spectrum of activity against all four classes of fungi. Strobilurin fungicides The strobilurin fungicides were developed recently. . each of which is in a unique class of chemistry. Azoxystrobin (Quadris) and kresoxim-methyl (Sovran. Most fungicides in this class are strongly bound to the cuticle (leaf surface) and so are not easily washed off the leaf. It has excellent activity against silver scurf of potato. They inhibit fungal respiration. Other strobilurins such as kresoxim-methyl and trifloxystrobin have only slight uptake by the leaf and are redistributed on the plant through vapor action. Sterol-inhibiting fungicides currently available in North Dakota (2000) include imazalil (several brand names). These fungicides are environmentally friendly. It is registered for use as a seed pelleting for sugarbeets for early season control of Aphanomyces seedling disease.Sterol inhibitor (ergosterol biosynthesis inhibitor) fungicides The sterol inhibitor fungicides inhibit the formation of ergosterol in the higher fungi. Cygnus) were the first of this class. registration expected in several years). Hymexazol (Tachigaren) is an isoxazole class of chemistry that is environmentally friendly and has excellent activity against Aphanomyces and also Pythium. New Classes of Fungicides Several other fungicides have been developed recently. and tetraconazole (registration pending for Eminent). as well as Rhizoctonia black scurf of potato and Fusarium dry rot. This is a methoxyacrylate class of chemistry which is related to naturally occurring products found in a group of forest mushrooms called "pine-cone mushrooms. Other strobilurins include trifloxystrobin (Flint). expected to be registered for the 2002 season) and a product with a related mode of action called famoxadone (Famoxate. triadimefon (Bayleton) triadimenol (Baytan). BAS 500 (Headline. may suppress spore formation in established infections. difenoconazole (Dividend seed treatment). triforine (Funginex. but not the water molds. tebuconazole (registered Raxil seed treatments. This prevents cell wall formation and stops growth of the fungus. propiconazole(Tilt). most are taken up by the plant (some are only locally systemic). moving into and up the leaf toward the tip. They have therapeutic activity. registration pending for Folicur foliar treatments).

and a pathogen that develops resistance to one of the benzimidazole fungicides will be resistant to all three. Repeated use of the same fungicide or fungicides with the same mode of action can lead to development of resistance. Fenhexamid (Elevate) is a hydroxyanilide class of chemistry that is used as a protectant fungicide. including activity against Sclerotinia (white mold). . dimethomorph (Acrobat) and propamocarb (Tattoo or Previcur) and the registration pending product zoxamide (Gavel). Cymoxanil is an acetimide fungicide that is locally systemic. The Fusarium dry rot pathogen and the silver scurf pathogen of potato have developed resistance to this class of fungicide across much of the United States and Canada. Resistance also can develop rapidly to the dicarboximides. requiring higher application rates to control Cercospora leafspot. has a short residual and has both curative and protectant properties. Resistance to the benzimidazoles and the phenylamides can occur rapidly. including many fields in Michigan. Resistance to Fungicides Fungi may develop resistance to fungicides. with resistant strains of fungi showing insensitivity to high rates of the fungicide. All four have good activity against the potato late blight pathogen and the downy mildews. Nevertheless. the sterol inhibitors. The Cercospora leafspot pathogen of sugarbeet is resistant to benzimidazole fungicides in many areas of the world. and the strobilurins. the Cercospora leafspot pathogen of sugarbeet has developed tolerance to triphenyltin hydroxide. Dimethomorph is a cinnamic acid derivative that is locally systemic and has curative and preventive activity. Fungi are more likely to develop resistance to systemic fungicides with a single specific mode of action. Zoxamide is an amide fungicide with a different mode of action from other potato late blight fungicides. Some fungicides that are in varying classes not listed above include the registered fungicides cymoxanil (Curzate). These three fungicides are closely related. This is called cross resistance. Fungi are less likely to develop resistance to the older protectant fungicides that have multiple sites of action against fungi. Some products such as acibenzolar (Actigard) and several others act to enhance the natural defenses of the plant. Minnesota and North Dakota. This tolerance is expressed as a reduced sensitivity to the fungicide. Currently there are no uses registered for North Dakota crops. Propamocarb is a carbamate fungicide that is systemic and is translocated into new foliage.Fluazinam (Omega) is a pyridinamide class of chemistry that is a contact fungicide with a broad spectrum of activity.

the sterol inhibitors. which reduces the likelihood that resistance will develop. Avoid reduced rates of fungicides. including the US8 genotype now present in North Dakota and many other parts of the United States and Canada. the dicarboximides and the strobilurins. This has been done with the phenylamide fungicides. This is a good strategy for resistance management of triphenyltin hydroxide. The gray mold fungus has rapidly developed resistance to the dicarboximide fungicides iprodione (Rovral) and vinclozolin (Ronilan) under greenhouse conditions.The potato late blight fungus can develop resistance rapidly to the phenylamides metalaxyl and mefenoxam. The presence of this new genotype has seriously limited the usefulness of this class of fungicide for potato late blight management. Apply a limited number of applications in a block at a critical period in the pathogen disease cycle. . Certain genotypes of the fungus have resistance. Certain grape pathogens have rapidly developed resistance to this class of fungicide in California. A different mode of action should be used at other less critical times in the disease cycle. These include: • • • • • • Tank mix with a fungicide with a different mode of action. so as to minimize the exposure of the "at risk" fungicide. Resistance Management There are several ways to retard the development of resistance. Do not use phenylamides as soil treatments against airborne pathogens. Use of these fungicides may be restricted to the most critical parts of the season. and resistance management practices are recommended for this class of fungicide. Cases of resistance to sterol inhibitors are fairly common. Limit the number of applications of an "at risk" fungicide per year. This has been recommended with some of the strobilurins. it is recognized that resistance can occur to this class of fungicide. Mancozeb or chlorothalonil can be tank mixed effectively with benzimidazole or phenylamide fungicides. Although the strobilurin fungicides have been used but a few years. and the strobilurins. and cases of resistance are already being reported world-wide. Alternate applications between or among two or more classes of fungicides with different modes of action. the sterol inhibitors. Although the sterol inhibitors and the phenylamides have some post-infection activity. they are best used in a preventive manner. These reduced rates may facilitate the development of resistance in fungi.

Most commonly used fungicides have a relatively low level of toxicity.sanitation is the first step to disease management for . and protection (fungicides). Growers need to integrate as many different management tools as possible for long term success. The main tools for management of dry bean rust are sanitation. long-sleeved shirt. especially for the maneb and mancozeb fungicides. When wheat is planted directly into standing wheat stubble. practical disease management usually involves several techniques. Tan spot of wheat can be reduced by sanitation. crop rotation. Management programs based on only one or two techniques can be effective in the short term but may become ineffective if used frequently. Stem rust of wheat is managed by host resistance. Homeowners should make every effort to remove diseased plant debris from the yard and garden in the fall . but a dry storage area is essential. Agri Tin) is an exception in that it is quite toxic.Use of Fungicides All pesticides are poisons. shoes. Many fungicides have information on the label restricting the number of days before harvest that the last application can be made to food crops. Care should be taken in storing liquid or flowable fungicides as some will not stand freezing. The development of fungicide resistance is a good example. Most fungicides can be stored successfully from year to year. Monitoring provides early warning for plant breeders of the buildup of new rust races and the need for alternative management for the grower. INTEGRATED DISEASE MANAGEMENT Effective. but triphenyltin hydroxide (Super Tin. the elimination of barberry (this prevents the sexual phase that produces new rust races). both sanitation and rotation are eliminated as management tools and growers must rely primarily on variety choice and fungicides for tan spot reduction. tolerant varieties and fungicides. researchers are developing varieties with improved tan spot resistance. All of these procedures must be done well and in a timely fashion to produce effective management. the epidemics of rust caused by newly prevalent races of the fungus is another. and by constant monitoring of the rust races by plant pathologists. Check the label for information on use and storage. and they should be used with care. Most have a 24-hour period during which re-entry in a field should be done only if wearing protective clothing (long pants. chemical-resistant gloves). crop rotation.

should be practiced. The disease forecasting information aids in predicting disease outbreaks and the need for fungicides. as this helps to anticipate disease-favoring weather and allows more timely preventive measures. exclusion. Finally. all the principles of management. eradication. use of resistant varieties. host resistance. Common causes of diseases.next spring. Meterologic etc. suffocatio n of roots in wet soil Frost. The use of these combined practices usually produces the most reliable and stable plant disease management. and protection. Some Disease Forecasting models for major crop diseases have been developed and are now available on the Internet or through toll-free telephone numbers. all growers should keep close watch on the weather forecasts. Whenever possible. How They Can Be Seen How They Magnificati Descriptio Reproduc Equipmen on Example n e t Required* s Deficiency -Visual (no -Nitrogen or excess special deficiency of essential equipment . zinc deficiency Deficiency or excess of water -Visual -Moisture stress or wilting. rotation. heat sterility Heat canker. Type Cause NonNutrition Infectiou s (Abiotic) Moisture Temperatur Cold or e heat Other Sun. -- Visual -- -- Visual -- . iron elements needed) deficiency . wind. Sanitation should be followed by seed treatment. Table 1. and timely fungicide applications.

bacterial blights of small grains.000host to microscop 100. leafspots. saline seeps). Bacterial blights of beans. Microscop 20-250 e (occasiona lly a hand lens) Bacteria Tiny single. Wheat streak mosaic. Cause Electron 20. large fruiting structures may develop from these filaments. -Visual -- sunscald. dry beans and sunflower) . smuts. lightning injury Salt injury (road salt. tiny Cell threadlike division filaments. fireblight.al Conditions Toxic Chemicals Salt. bacterial wilt of cucumber . ozone injury Rusts. etc.000 e Viruses Very tiny rod-shaped or spherical particles. tree cankers.000 manufactu e re virus . air pollutants. apple scab. powdery mildew. Microscop 400-1.Cell celled division organisms. wheat and barley scab. white mold (canola. barley Infectiou Fungi s (Biotic) Grow as Spores.

000 inch in size when magnified 100 times would appear to be 1/10 inch in size. Nematodes Tiny Eggs Roundwor ms No serious problem in North Dakota. a pathogen 1/1. Soybean cyst nematode and pine wilt nematode occur in neighbori ng states. Phytoplasm Very tiny Division as organisms without a cell wall. For example. the free encyclopedia Jump to: navigation. tobacco mosaic.composed of RNA with a protein coat.000. and Latin remedium (restoring balance or remediation) describes the treatment of environmental problems . plant).000 inch in size when magnified 100. February 2001 Microscop 1-60 e (naked eye for larger forms) Phytoremediation From Wikipedia. *The number of times a pathogen must be magnified to be visible.000 times would appear to be 1/10 inch in size. stripe mosaic. cucumber mosaic. Electron 20. no definite shape. EB-31. a pathogen 1/1. potato viruses. Revised. search Phytoremediation (from the Ancient Greek φυτο (phyto.000 e Aster yellows (purple top in potato and tomato).000microscop 50.

Examples where phytoremediation has been used successfully include the restoration of abandoned metalmine workings. Phytoremediation refers to the natural ability of certain plants called hyperaccumulators to bioaccumulate. or eliminate metals.3 Phytotransformation 4 The role of genetics 5 Hyperaccumulators and biotic interactions o 5.or render harmless contaminants in soils. or air. pesticides. this technology has become . pesticides. Many plants such as mustard plants. Phytoremediation consists of mitigating pollutant concentrations in contaminated soils.1 Table of hyperaccumulators 6 See also 7 References 8 Bibliography 9 External links [edit] Application Phytoremediation may be applied wherever the soil or static water environment has become polluted or is suffering ongoing chronic pollution. have been mitigated in phytoremediation projects worldwide. Over the past 20 years. as opposed to mechanical cleanup methods such as soil excavation or pumping polluted groundwater. reducing the impact of sites where polychlorinated biphenyls have been dumped during manufacture and mitigation of on-going coal mine discharges.2 Phytostabilization o 3. cost-effective and non-environmentally disruptive technology. solvents. Phytoremediation is considered a clean. or air. degrade. Contaminants such as metals. Contents [hide] • • • • • • • • • 1 Application 2 Advantages and limitations 3 Various phytoremediation processes o 3. water.(bioremediation) through the use of plants that mitigate the environmental problem without the need to excavate the contaminant material and dispose of it elsewhere. solvents. and crude oil and its derivatives. explosives. degrade. explosives[1]. water. with plants able to contain. crude oil and its derivatives. and various other contaminants from the media that contain them. alpine pennycress and pigweed have proven to be successful at hyperaccumulating contaminants at toxic waste sites.1 Phytoextraction o 3.

uranium. tolerance to toxicity.increasingly popular and has been employed at sites with soils contaminated with lead. especially metals. which in itself does not resolve the problem of contamination) o the survival of the plants is affected by the toxicity of the contaminated land and the general condition of the soil. Phytostimulation — enhancement of soil microbial activity for the degradation of contaminants. This process is . or immobilization (phytostabilization). from primary level consumers upwards or requires the safe disposal of the affected plant material. into the plants which then pass into the food chain. Limitations: o phytoremediation is limited to the surface area and depth occupied by the roots. However. for example. and arsenic. one major disadvantage of phytoremediation is that it requires a long-term commitment. it is not possible to completely prevent the leaching of contaminants into the groundwater (without the complete removal of the contaminated ground. degradation (phytodegradation). often resulting in their inactivation. Phytotransformation — chemical modification of environmental substances as a direct result of plant metabolism. • [edit] Various phytoremediation processes A range of processes mediated by plants or algae are useful in treating environmental problems: • • • • Phytoextraction — uptake and concentration of substances from the environment into the plant biomass. as the process is dependent on plant growth. [edit] Advantages and limitations • Advantages: o the cost of the phytoremediation is lower than that of traditional processes both in situ and ex situ o the plants can be easily monitored o the possibility of the recovery and re-use of valuable metals (by companies specializing in “phyto mining”) o it is potentially the least harmful method because it uses naturally occurring organisms and preserves the environment in a more natural state. typically by organisms that associate with roots. o slow growth and low biomass require a long-term commitment o with plant-based systems of remediation. and bioaccumulation capacity. by limiting the leaching of substances from the soil. o bio-accumulation of contaminants. Phytostabilization — reducing the mobility of substances in the environment.

Advantages: The main advantage of phytoextraction is environmental friendliness. or phytomining. Phytostimulation can also involve aquatic plants supporting active populations of microbial degraders. After harvest. In general.[2] Phytovolatilization — removal of substances from soil or water with release into the air. Traditional methods that are used for cleaning up heavy metal-contaminated soil disrupt soil structure and reduce soil productivity. The pollutants remain absorbed in or adsorbed to the roots. The plants absorb contaminants through the root system and store them in the root biomass and/or transport them up into the stems and/or leaves. so the growth/harvest cycle must usually be repeated through several crops to achieve a significant cleanup. sometimes as a result of phytotransformation to more volatile and/or less polluting substances. the cleaned soil can support other vegetation. After the process. Rhizofiltration — filtering water through a mass of roots to remove toxic substances or excess nutrients. and induced or assisted hyper-accumulation. A living plant may continue to absorb contaminants until it is harvested. in which a conditioning fluid containing a chelator or another agent is added to soil to increase metal solubility or mobilization so that the plants can absorb them more easily. In many cases natural hyperaccumulators are metallophyte plants that can tolerate and incorporate high levels of toxic metals. where plants naturally take up the contaminants in soil unassisted. Disadvantages: As this process is controlled by plants. Two versions of phytoextraction: • • natural hyper-accumulation. Phytoextraction has been growing rapidly in popularity worldwide for the last twenty years or so. At the time of disposal. it takes more time than anthropogenic soil clean-up methods. whereas phytoextraction can clean up the soil without causing any kind of harm to soil quality. as in the stimulation of atrazine degradation by hornwort. this process has been tried more often for extracting heavy metals than for organics. is also being experimented with. 'Mining with plants'. Another benefit of phytoextraction is that it is less expensive than any other clean-up process. [edit] Phytoextraction Phytoextraction (or phytoaccumulation) uses plants or algae to remove contaminants from soils. sediments or water into harvestable plant biomass (organisms that take largerthan-normal amounts of contaminants from the soil are called hyperaccumulators). .• • also known as rhizosphere degradation. contaminants are typically concentrated in the much smaller volume of the plant matter than in the initially contaminated soil or sediment. a lower level of the contaminant will remain in the soil.

Pollutants become less bioavailable. or the Chinese Brake fern (Pteris vittata). certain plants. which sequester lead in their biomass. using Willow (Salix viminalis): In 1999. a hyperaccumulator of these metals at levels that would be toxic to many plants. explosives. An example application of this sort is using a vegetative cap to stabilize and contain mine tailings. and human exposure is reduced. phytostabilization focuses mainly on sequestering pollutants in soil near the roots but not in plant tissues.[6] [edit] Phytotransformation In the case of organic pollutants. microorganisms living in association with plant roots may metabolize these substances in soil or water. For example. Zinc (Zn). as willow has some specific characteristics like high transport capacity of heavy metals from root to shoot and huge amount of biomass production. or Poplar trees. Hemp Dogbane (Apocynum cannabinum). and other xenobiotic substances. Salt-tolerant (moderately halophytic) barley and/or sugar beets are commonly used for the extraction of Sodium chloride (common salt) to reclaim fields that were previously flooded by sea water. using Alpine pennycress (Thlaspi caerulescens). selenium and organic pollutants such as polychlorinated biphenyls (PCBs) have been removed from soils by transgenic plants containing genes for bacterial enzymes. immobilize the pollutants by adsorption or accumulation. one research experiment performed by Maria Greger and Tommy Landberg suggested Willow (Salix viminlais) has a significant potential as a phytoextractor of Cadmium (Cd). such as pesticides. In other cases.[4] Mercury. such as Cannas. Lead. the plant's presence can reduce wind erosion. These . and livestock. render these substances non-toxic by their metabolism.[3] Cadmium and zinc. Ragweed (Ambrosia artemisiifolia). solvents. or the plant's roots can prevent water erosion. using the Sunflower (Helianthus annuus). Chinese Brake fern stores arsenic in its leaves. using Indian Mustard (Brassica juncea). can be used also for production of bio energy in the biomass energy power plant. wildlife.[5] • • • • [edit] Phytostabilization Phytostabilization focuses on long-term stabilization and containment of the pollutant. and provide a zone around the roots where the pollutant can precipitate and stabilize. Caesium-137 and strontium-90 were removed from a pond using sunflowers after the Chernobyl accident. and Copper (Cu). On the other hand.Examples of phytoextraction (see also 'Table of hyperaccumulators'): • • • Arsenic. Cadmium. the presence of copper seems to impair its growth (see table for reference). Unlike phytoextraction. industrial chemicals. a hyperaccumulator.

complex and recalcitrant compounds cannot be broken down to basic molecules (water.[7] After uptake of the xenobiotics. plant biomolecules such as glucose and amino acids are added to the polarized xenobiotic to further increase the polarity (known as conjugation). In the final stage of phytotransformation (Phase III metabolism). This is again similar to the processes occurring in the human liver where glucuronidation (addition of glucose molecules by the UGT (e. The increased polarity also allows for easy transport of the xenobiotic along aqueous channels. This is known as Phase I metabolism. carbon-dioxide. In the second stage of phytotransformation. Genes for phytoremediation may originate from a micro-organism or may be transferred from one plant to another variety better adapted to the environmental conditions at the cleanup site. UGT1A1) class of enzymes) and glutathione addition reactions occur on reactive centres of the xenobiotic. the plants reduce toxicity (with exceptions) and sequester the xenobiotics in phytotransformation. etc. and. . However. Whereas in the human liver enzymes such as Cytochrome P450s are responsible for the initial reactions. The term "Green Liver Model" is used to describe phytotransformation. a sequestration of the xenobiotic occurs within the plant.[9] Researchers have also discovered a mechanism in plants that allows them to grow even when the pollution concentration in the soil is lethal for non-treated plants. in plants enzymes such as nitroreductases carry out the same role. Phase I and II reactions serve to increase the polarity and reduce the toxicity of the compounds. This ensures that the xenobiotic is safely stored. plants involved in phytotransformation may need to be maintained in a closed enclosure. and does not affect the functioning of the plant. as plants behave analogously to the human liver when dealing with these xenobiotic compounds(foreign compound/pollutant). or for introducing new capabilities into plants. genes encoding a nitroreductase from a bacterium were inserted into tobacco and showed faster removal of TNT and enhanced resistance to the toxic effects of TNT.) by plant molecules. Trinitrotoluene phytotransformation has been extensively researched and a transformation pathway has been proposed. The xenobiotics polymerize in a lignin-like manner and develop a complex structure that is sequestered in the plant. and. similar to the way that the human liver increases the polarity of drugs and foreign compounds (Drug Metabolism). Hence.[8] [edit] The role of genetics Breeding programs and genetic engineering are powerful methods for enhancing natural phytoremediation capabilities. hence. the term phytotransformation represents a change in chemical structure without complete breakdown of the compound. For example.g. hence. known as Phase II metabolism. preliminary studies have shown that these plants can be toxic to small animals (such as snails). plant enzymes increase the polarity of the xenobiotics by adding functional groups such as hydroxyl groups (-OH). although many exceptions to the rule are seen.

cobalt. [edit] Table of hyperaccumulators Plant Nutrient Deficiencies Identifying Plant Problems By Marie Iannotti. or more than 10.com Gardening Ads • • Gardening Leaves . such as exogenous polyamines.cmtevents. allow the plants to tolerate concentrations of pollutants 500 times higher than untreated plants.com 2nd CommercialFarm AfricaNew Structures Of Agri-Investment 31 Jan to 1 Feb 2012. mutualism (including mycorrhizae.com Guide See More About: • • • plant problems fertilizer garden maintenance Sponsored Links The Humic ExpertsHigh quality humates and humic acid products for healthy plants & soilhumates.com SPAD Chlorophyll MeterImprove Your Crop Quality & Yield Manage Your Nitrogen Managementwww. chromium or lead. Dubai UAEwww. pollen and seed dispersal). and to absorb more pollutants. About. biodegradable compounds. copper. including protection. [edit] Hyperaccumulators and biotic interactions A plant is said to be a hyperaccumulator if it can concentrate the pollutants in a minimum percentage which varies according to the pollutant involved (for example: more than 1000 mg/kg of dry weight for nickel.[10] This capacity for accumulation is due to hypertolerance. interferences with neighbour plants of different species. or phytotolerance: the result of adaptative evolution from the plants to hostile environments through many generations.Some natural. A number of interactions may be affected by metal hyperaccumulation.000 mg/kg for zinc or manganese). and biofilm. commensalism.konicaminolta.

Nutrients that are needed in relatively large amounts are called the macronutrients. but in much smaller quantities. molybdenum and zinc. Unfortunately many problems have similar symptoms and sometimes it is a combination of problems. the plant will not be able to take in nutrients no matter how rich your soil may be. Be sure you eliminate the obvious before you kill your plants with kindness. For a definitive diagnoses. even when the soil is wet.• • • How to Plant a Garden Flower Garden Gardening Plant Pests Sponsored Links Bulk Chelated MineralsChelated minerals in bulk wholesale Wholesale prices worldwide deliverywww.com Not all plant problems are caused by insects or diseases. There are a handful of additional nutrients that are required for plant growth. The following chart outlines some possible problems. sulfur and magnesium. potassium. Extreme cold or heat will slow plant growth and effect flowering and fruit set.vgdusa. manganese. A second requirement is the appropriate soil pH for the plant being grown. copper. So one requirement of sufficient plant nutrition is water. Plant nutrient deficiencies often manifest as foliage discoloration or distortion.stockton-agrimor. Some plants are fussier than others.com Stockton AgrimorA new generation of biofungicides Plant disease control . contact your local cooperative extension service. • • • • Check first for signs of insects or disease. but if the soil pH is too acidic or alkaline. Each plant prefers a specific pH range to be able to access the nutrients in the soil. These micronutrients include: boron. Too much fertilizer can result in salt injury. Plants require a mix of nutrients to remain healthy. Water transfers the nutrients from the soil to the plant roots. Your plants may look scorched or they may wilt. Foliage discoloration and stunted plants can easily be caused by soil that is too wet and drains poorly or soil that is too compacted for good root growth. iron. phosphorus. calcium. Sometimes an unhealthy plant is suffering from a nutrient deficiency or even too much of any one nutrient. .No residueswww. Plant macronutrients include: nitrogen. All of these nutrients are taken in through the roots.

Notes: Many forms of nitrogen are water soluble and wash away. such as Epson Salts. Phosphorus (P) o o o Symptoms: Small leaves that may take on a reddish-purple tint. Also gypsum. Magnesium (Mg) o o Symptoms: Slow growth and leaves turn pale yellow. 'ammonium' or 'urea'. generally at the bottom of the plant. Sources: Any compound containing the word 'calcium'. Interveinal chlorosis (yellowing between the leaf veins) develops. Reduced fruit or seed production. Leaf tips can look burnt and older leaves become almost black. Sources: Compounds containing the words 'phosphate' or 'bone'. sometimes just on the outer edges. tip burn of cabbage and brown/black heart of escarole & celery. The growing tip may die. Notes: Very dependent on pH range. Remaining foliage is often light green. Also manure. Contributes to blossom end rot in tomatoes.Plant Nutrient Deficiency Symptoms Macronutrients Calcium (Ca) o o o Symptoms: New leaves are distorted or hook shaped. Stems may also yellow and may become spindly. Also greensand. Growth slows. Nitrogen (N) o o o Symptoms: Older leaves. New growth may be yellow with dark spots. Sources: Any compound containing the words: 'nitrate'. will yellow. . Potassium (K) o Symptoms: Older leaves may look scorched around the edges and/or wilted. Sources: Compounds containing the word 'magnesium'. Notes: Not often a deficiency problem and too much will inhibit other nutrients.

Stunts growth. Seed stalks also become limp and bend over. Failure to bloom. shoots and fruit diminished in size. Copper (Cu) o o Symptoms: Stunted growth. remaining foliage turns light green. Terminal (end) buds may die. Leaves can become limp. Sources: Compounds containing the words 'borax' or 'borate'. Notes: More prevalent in dry weather. Sources: Compounds containing the word 'sulfate'. Notes: Sometimes confused with nitrogen deficiency. Sources: Compounds containing the words 'copper'. Micronutrients Boron (B) o o Symptoms: Poor stem and root growth. Sources: Compounds containing the words 'molybdate' or 'molybdic'. Leaves. Sulfur (S) o o o Symptoms: New growth turns pale yellow. May develop dark or dead spots. Sources: Compounds containing the words 'manganese' or 'manganous' Molybdenum (Mo) o o o Symptoms: Older leaves yellow. often starting between veins. Leaves can become narrow and distorted. Zinc (Zn) . older growth stays green. 'cupric' or 'cuprous'.o Sources: Compounds containing the words 'potassium' or 'potash'. or drop. Younger leaves turn pale yellow. Manganese (Mn) o o Symptoms: Growth slows. curl. Witches brooms sometimes form.

so there is not a one-to-one relationship between the molarity and the osmolarity of a . However. For the osmole unit. such as salts. osmolality 5 See also 6 References [edit] Types of solutes Osmolarity is distinct from molarity because it measures moles of solute particles rather than moles of solute. Whereas molarity measures the number of moles of solute per unit volume of solution. Molarity and osmolarity are not commonly used in osmometry because they are temperature dependent. tonicity 4 Plasma osmolarity vs. The osmolarity of a solution is usually expressed as Osm/L (pronounced "osmolar"). if the concentration of solutes is very low.[1] Ionic compounds. Contents [hide] • • • • • • 1 Types of solutes 2 Definition 3 Osmolarity vs.o o o Symptoms: Yellowing between veins of new growth. Osmolarity From Wikipedia. The distinction arises because some compounds can dissociate in solution. defined as the number of osmoles (Osm) of solute per litre (L) of solution (osmol/L or Osm/L). see Osmole (unit). osmolarity measures the number of osmoles of solute particles per unit volume of solution. Terminal (end) leaves may form a rosette. Notes: Can become limited in higher pH. search This article is about osmolarity. osmolarity and osmolality are considered equivalent. Osmolarity is the measure of solute concentration. whereas others cannot. the free encyclopedia Jump to: navigation.[1] Osmolality is a measure of the osmoles of solute per kilogram of solvent (osmol/kg or Osm/kg). in the same way that the molarity of a solution is expressed as "M" (pronounced "molar"). can dissociate in solution into their constituent ions. This is because water changes its volume with temperature (See: Vapour pressure of water). Sources: Compounds containing the word 'zinc'.

Osmolality can be measured using an osmometer which measures colligative properties. or 1 mole glucose + 1 mole NaCl. Vapor pressure. hypertonic. electrostatic effects cause φ to be smaller than 1 even if 100% dissociation occurs (see Debye-Hückel equation). hyperosmotic. but different. for sucrose). For example: glucose has n of 1. C is the molar concentration of the solute. In the simplest case it is the degree of dissociation of the solute. a 1 M NaCl solution is a 2 Osm NaCl solution). such as Freezing-point depression.5 mole NaCl.[1] Multiple compounds may contribute to the osmolarity of a solution. the index i represents the identity of a particular solute. hyposmotic) are not synonymous with the terms ending in -tonic (isotonic. [edit] Osmolarity vs.. whereas tonicity takes into account the total concentration of only non-penetrating solutes. while NaCl has n of 2.e. a 3 Osm solution might consist of: 3 moles glucose.g. φ is between 0 and 1 where 1 indicates 100% dissociation. which accounts for the degree of non-ideality of the solution. For salts. Then. there are 2 osmoles of solute particles (i.solution. For example. ions) into which a molecule dissociates. the terms ending in -osmotic (isosmotic. For example. For example. However. and form only 1 osmole of solute per 1 mole of solute.g.[1] [edit] Definition The osmolarity of a solution can be calculated from the following expression: where • • • • φ is the osmotic coefficient.[1] . for every 1 mole of NaCl in solution.5 moles NaCl. tonicity Osmolarity and tonicity are related. Both sodium and chloride ions affect the osmotic pressure of the solution. or Boiling-point elevation. concepts.[1] Nonionic compounds do not dissociate. or 2 moles glucose + 0.ions. hypotonic). The terms are related in that they both compare the solute concentrations of two solutions separated by a membrane. Thus. φ can also be larger than 1 (e. Thus. n is the number of particles (e. The terms are different because osmolarity takes into account the total concentration of penetrating solutes and non-penetrating solutes. or 1. sodium chloride (NaCl) dissociates into Na+ and Cl. a 1 M solution of glucose is 1 Osm. or any other such combination.

Non-penetrating solutes cannot cross the cell membrane.if the total concentration of solutes in one compartment is different from that of the other. drawing water with it and thus causing no net change in solution volume. and therefore osmosis of water must occur for the solutions to reach equilibrium. Using a gas-permeable closure. In more simple terms. [edit] Plasma osmolarity vs. A solution can be both hyperosmotic and isotonic. osmolality is an expression of solute osmotic concentration per mass. Paffen. V. an inhibitor of . Addition of silverthiosulphate (STS).Penetrating solutes can diffuse through the cell membrane. According to IUPAC. whereas osmolarity is per volume of solvent (thus the conversion by multiplying with the mass density).025 g/ml for blood plasma. osmolality is the quotient of the negative natural logarithm of the rational activity of water and the molar mass of water. but one of the ions cannot cross the membrane. whereas osmolarity is the product of the osmolality and the mass density of water (also known as osmotic concentration).[3] ca is the (anhydrous) solute concentration in g/ml . causing momentary changes in cell volume as the solutes "pull" water molecules with them. the intracellular fluid and extracellular can be hyperosmotic.03 g/ml.[1] For example. but isotonic . plasma osmolarity is 1-2%[2] less than osmolality. Klerk Abstract: The effects of different growth regulators and inhibitors on sprouting and outgrowth of isolated axillary buds of rose and apple in vitro were evaluated. Telgen. ROLE OF PLANT HORMONES IN LATERAL BUD GROWTH OF ROSE AND APPLE IN VITRO Authors: H. A. suggesting an inhibiting effect of accumulating ethylene. Elagöz. The response was dependent on the original position of the axillary bud on the main stem. Mil.not to be confused with the density of dried plasma Since ca is slightly larger than 0. Budbreak in ‘Madelon’ rose was stimulated by benzyladenine (BA). osmolality Plasma osmolarity can be calculated from plasma osmolality by the following equation:[2] Osmolarity = osmolality * (ρsol .ca) Where: • • ρsol is the density of the solution in g/ml. G. which is 1. A. shoot length was increased in ‘Madelon’ but not in ‘Motrea’ rose.

In apple. increased shoot length significantly in both rose varieties and in apple. the presence of ACC decreased the multiplication rate.ethylene action. 312512 bytes) Citation . • • Full Text (PDF format. the stimulating effect of STS depended upon the auxin concentration in the medium. Inhibiting ethylene biosynthesis with aminoethoxyvinylglycine (AVG) or stimulating it with 1-aminocyclopropane-1-carboxylic acid (ACC) had no effect on average shoot length in rose. Only in ‘Madelon’. Abscisic acid (ABA) reduced or inhibited BA-induced prouting of isolated axillary buds of ‘Madelon’ rose.