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Mineral Nutrition and Ion Transport

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    CHAPTER 7:

    Mineral nutrition and ion transport

    Section "A"

    Introduction

    The supply and absorption of specific chemical compounds needed for normal growth and

    metabolism of plants is defined asnutrition while the chemical compounds that functions as

    raw material for synthesis of different structural and functional substance of plants are

    termednutrients. The inorganic material obtained from soil which are used as raw material

    by plants are calledmineral nutrients. Absorption utilization and assimilation of inorganic

    compound or minerals by plants for synthesis of essential material for their growth,

    development, structure and physiology is called mineral nutrition.

    Essential elements The mineral nutritional elements are available to the plant as ion presentin soil water absorbed through root. Nutrient elements may occur in the soil (a) in the aueous

    solution, (b) adsorbed on organic or inorganic soil colloids, (c) in the form of an insoluble

    inorganic compound, and (d) as a constituent of organic compounds, either as a residue of

    plants or animals or in living organisms. The upta!e of nutrients by the plant roots is closely

    related to the form in which the elements occur. Many factors influence nutrient upta!e for

    plants. "ons can be readily available to roots or could be #tied up# by other elements or the

    soil itself. $oil too high in p% (al!aline) or too low (acid) ma!es minerals unavailable to

    plants.

    Liebig (!#$proposed the law of minimum which states that productivity of a soil is

    dependent upon the proportionate occurrence of deficient mineral.

    %ulius &on Sac's( !#$)&erman botanist, demonstrated, for the first time, that plants could

    be grown from seedling to maturity in a defined nutrient solution in complete absence of soil.

    This techniue of growing plants in a nutrient solution is !nown as '*dro+onics.

    Arnon and Stout (,-,$physiologists have given three criteria for plant nutrient

    essentiality '

    . $ome elements are essential for supporting normal growth and reproduction. "n the

    absence of the element the plants do not complete their life cycle or set the seeds.

    *. The reuirement of the element must be specific and the element cannot be replaced bythe presence of another element. "n other words, deficiency of any one element cannot be

    met by supplying some other element.

    +. The element must be directly involved in the metabolism of the plant.

    Two criteria by pstein, *--'

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    i. The element is part of a molecule that is an intrinsic component of the structure or

    metabolism of a plant.

    ii. The plant can be so severely deprived of element that it e/hibits abnormalities in itsgrowth, development or reproduction that is, its 0performance1 in comparison with

    plants not so deprived.

    2or e/ample, agnesium is a constituent of the chlorophyll molecule and is essential for

    photosynthesis, so it cannot be replaced by any other element for the same function . "t is also

    reuired as aco/actorby many enzymes involved in cellular res+iration and metabolic

    +at' 0a*s.

    Si1teen c'emical elements are important in which carbon) '*drogen) and o1*genare

    supplied by air and water, which comprise about 345 of the fresh weight of the plant, at least

    + other chemical elements, called the essential inorganic nutrients, are needed for plant6s

    growth and survival. The si/teen chemical elements are divided into two main groups'

    a. 2on3ineral 2utrients'The NonMineral Nutrients are'*drogen (H$) o1*gen (4$)and carbon (C$."n a process called +'otos*nt'esis)plants use energy from the sun tochangecarbon dio1ide (C45 3 carbon and o1*gen$ and 0ater (H543 '*drogen and

    o1*gen$into starches and sugars.

    b. ineral + other chemical elements,which come from the soil, are dissolved in water

    and absorbed through a plant6s roots called the essential inorganic nutrients, are neededfor plant growth.

    7n the basis of average concentration in plants , Hoagland (,$ divided essential elements

    into two categories'

    . acronutrients:Macronutrients are generally present in plant tissues in largeconcentrations o/ .#3#.# mg +er gram o/ dr* matter.The macronutrientsincludeCarbon (C$ ) '*drogen (H$) o1*gen(4$) nitrogen)(2$ +'os+'orous(P$)

    sul+'ur(S$) +otassium (6$) Calcium (Ca$and magnesium(g$.

    *. 2itrogen) Potassium) and P'os+'orousare obtained from the soil and are the +rimar*macronutrients.Calcium) agnesium) and Sul+'urare the secondar*

    macronutrientsneeded in lesser uantity.

    +. icronutrients or trace elements:Micronutrients are present in very small amounts inplant body in concentration ofeual or less t'an #. mg +er gram o/ dr* matter.These

    includeiron (8e$) manganese (n$ ) Co++er (Cu$) mol*bdenum (o$) 9inc (n$)

    boron) (;$ c'lorine (Cl$ .Many of them arecom+onents or co/actors o/

    en9*mes.$ome are essential for electron trans/er.

    There are some beneficial elements such as sodium) silicon) cobalt and selenium) in addition

    to the 8 essential elements named above They are reuired by higher plants.

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    ssential elements can also be grouped into following types on the basis of their diverse

    functions '

    i. 8rame0or< elements:3 Carbon (C $) H*drogen (H$) 41*gen (4$synthesize the

    cellulose and other components of cell walls. 9alcium (9a) is component of middle

    lamella.

    ii. Proto+lasmic elements :3 Carbon (C$ ) '*drogen (H$) o1*gen(4$) nitrogen)(2$+'os+'orous(P$) sul+'ur(S$) iron (8e$and magnesium(g$give rise to protoplasmic

    constituents li!e proteins, nucleic acids, hormones, vitamins, phospholipids,,photosynthetic pigments, etc.

    iii. 4smotic +otential :3$ome essential elements, e. g., nitrate, potassium, sulphate etc. can

    alter the osmotic potential of a cell.Potassiumplays an important role in the opening and

    closing of stomata and turgor movements in various plants.

    iv. Energ* transducers :3agnesium occurs in c'loro+'*ll and +'os+'orous in ATP areessential elements that are components of energyrelated chemical compounds in plants,

    converting one form of energy into another.

    v. En9*matic e//ects:3$ome elements that acti&ate or in'ibit en9*mes) e.g.

    6)Ca)g)n)n)Cl)2i)Cu) etc. g5=is an activator for both ribulose bisphosphate

    carbo/ylaseo/ygenase and phosphoenol pyruvate carbo/ylase, both of which are critical

    enzymes in photosynthetic carbon fi/ation,n5=is an activator of alcohol

    dehydrogenase and oof nitrogenase during nitrogen metabolism.

    vi. ;alancing elements :3 Calcium ) magnesium +otassium counteract the to/ic effect of

    other minerals by ion balancing.

    vii. 8e) n) Cu) n and o are function as co enzymes

    Section ";"

    Role o/ acro3 and icro3nutrients and T'eir >e/icienc* s*m+toms

    .

    2ame o/

    elements

    Element

    obtained in t'e/orm o/

    Regions o/

    +lant 0'ere

    element

    reuired

    8unctions >e/icienc* s*m+toms

    Macronutrient

    s

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    Nitrogen (N)

    Nitrogen cane/ist in a soil

    organic form, or

    as anammonium ion

    (N%:;)nitrite

    ion (N7*),ornitrate ion

    (N7+).

    Nitrogen isreuired by all

    parts of a

    plant,particularly the

    meristematic

    tissues and themetabolically

    active cells.

    Nitrogen is one of

    the mahosphorus

    (>)

    >hosphorus is

    absorbed by the

    plants from soilin the form of

    phosphate ionsas %*>7:

    orthophosphate.

    >hosphate is

    easilyredistributed in

    most plants

    from one organto another and

    is lost from

    older leaves,

    accumulatingin younger

    leaves,developing

    flowers and

    seeds.The

    meristemregion of

    growing plants

    is high inphosphorus.

    >hosphorus is aconstituent of cell

    membranes, certain

    proteins, all nucleicacids and

    nucleotides,

    involved in energytransfer such as

    AT> and A?>. "t

    activates

    coenzymes foramino acid

    production used in

    proteinsynthesis@and it is

    involved in many

    other metabolicprocesses reuired

    for normal growth,

    such as

    photosynthesis,

    glycolysis,respiration, and

    fatty acid synthesis.

    $tunted growth, dar!

    green leaves with aleathery te/ture, and

    reddish purple leaf tips

    and margins.Maturity isoften delayed.

    >otassium () "t is absorbed as

    potassium ion(;).

    =euired in

    more abundantuantities in

    the

    %elps to determine

    an anioncationbalance in cells and

    activates enzymes

    $mall plants, brown

    margins on lower leaves,small wea! stems,

    lodging of plants, poor

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    meristematic

    tissues, buds,

    leaves and root

    tips.

    to metabolize

    carbohydrates for

    the manufacture of

    amino acids andproteins, opening

    and closing of

    stomata, activationof enzymes and in

    the maintenance of

    the turgidity ofcells, facilitates cell

    division and growth

    by helping to move

    starches and sugarsbetween plant

    parts,disease

    resistance.

    yield and uality.As in Nand >, ; is easily

    redistributed from mature

    to younger organs, so

    symptoms first appear inolder leaves.

    Magnesium

    (Mg)

    "t is absorbed as

    magnesium ion

    (Mg*;).

    "t is used forfruit and nut

    formation and

    essential forgermination of

    seeds. Beaves'

    withdrawn

    from ageingleaves and

    e/ported to

    developingseeds

    is a critical

    structuralcomponent of the

    chlorophyll

    molecule and isnecessary for

    functioning of plant

    enzymes to produce

    carbohydrates,sugars and fats."t is

    an enzyme activator

    in the synthesis ofnucleic acids (?NA

    and =NA).

    /tensiveinterveinalc'lorosiswhic

    h starts with basal leaves

    and progresses to younger

    leaves (it is mobile).

    9alcium

    (9a*;)

    "t is absorbed as

    calcium ion

    (9a*;).Adsorbe

    d 9a*; is

    important forsoil structure by

    promoting theaggregation of

    soil particles.

    This improveswater and root

    penetration

    Meristematic

    and

    differentiating

    tissues. "t is

    immobile(non

    translocatable)within plants

    and remains in

    the older tissuethroughout the

    growing

    9alcium is involved

    in many plant

    processes, including

    cell elongation, cell

    division,germination, pollen

    growth, activatesenzymes,and is a

    structural

    component of cellwalls, . 7ne of its

    most important

    "nhibition of bud growth,

    death of root tips,

    cupping of mature leaves,

    wea! growth.,

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    through the soil

    and maintains

    the stability ofsoil particles.

    season.9alciu

    m is easily

    leached.

    functions is the

    maintenance of

    membrane

    permeability andcell integrity.

    $ulphur ($)

    $ulfate ($7:C)

    from the soil isthe primary

    source of

    $.$ulfur is also

    ta!en up byleaves in

    gaseous form as$7*.

    $tem and roottips young

    leaves@

    remobilizedduring

    senescence

    (the growth

    phase in aplant or plant

    part as a leaffrom fullmaturity to

    death).

    "t is a structuralcomponent of

    amino acids,

    proteins, vitamins

    and enzymes and isessential to produce

    chlorophyll.

    &eneral chlorosis of leaf,

    including vascularbundles.$ulfur is not

    easily mobilized within

    theplant.Therefore,chlorosis,

    occur in young tissues

    before older ones.

    Micronutrient

    Dinc Dn

    (immobile)

    Most zinc in soilis found in

    different

    minerals withonly a smallpercentage

    being adsorbed

    in ionic form onsoil and organic

    matter e/change

    sites.

    verywhere

    "t activates various

    enzymes especially

    carbo/ylases, partof carbonicanhydrase and

    various

    dehydrogenases,needed for au/in

    synthesis.

    "nterveinal chlorosis of

    the upper (youngest)leaves. Afterwards, shootgrowth slows down,

    giving the affected plant

    parts a rosetteli!eappearance.

    Manganese

    Mn

    ("mmobile)

    The most

    important form

    for upta!e byroots is Mn;;,

    but it is also

    commonlyfound as o/ides

    of Mn;;; and

    Mn ;;;;(Mn*7+,

    Beaves and

    seeds

    Manganese in the

    plant participates in

    several importantprocesses including

    photosynthesis and

    nitrogen andcarbohydrate

    metabolism.

    The absence of

    Manganese causes

    disorganization ofchloroplast thyla!oid

    membranes. 9hlorosis

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    Mn7*, etc.).

    "ron, 2e

    ("mmobile)

    >lants obtain

    iron in the formof ferric ions

    (2e+;). "t is

    reuired inlarger amounts

    as compare to

    other

    micronutrients.

    verywhere ,

    collects alongleaf veins.

    "mportant

    constituent of

    proteins involved inthe transfer of

    electrons li!eferredo/in and

    cytochromes,

    reversibly o/idisedfrom 2e*; to 2e+;

    during electron

    transfer,activates

    catalase enzyme,reuired for

    synthesis ofchlorophyll.

    /tensive interveinal

    chlorosis, starting with

    younger leaves (iron isrelatively immobile).

    9opper (9u)

    "t is absorbed as

    cupric ions

    (9u*;).

    verywhere

    ssential for the

    overall metabolismin plants.More than

    half of the copper is

    located in thechloroplasts and

    participates in

    photosynthetic

    reactions. "t is alsofound in other

    enzymes involved

    with protein andcarbohydrate

    metabolism.

    ?iebac! of shoots

    Molybdenum

    (Mo)

    >lants obtain it

    in the form of

    molybdate ions.

    verywhere,

    the synthesis

    of proteins is

    bloc!ed andplant growth

    ceases. =oot

    nodule(nitrogen

    fi/ing) bacteria

    also reuire it.$eeds may not

    ssential

    component of two

    enzymes involved

    with nitrogenmetabolism.

    >ale green leaves with

    rolled or cupped margins.

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    form

    completely,

    and nitrogen

    deficiency mayoccur if plants

    are lac!ing

    molybdenum.

    Eoron (E)

    The soluble

    form isprimarily boric

    acid (E(7%)+).

    "n neutral to

    acid soils, boricacid has no

    charge and can,therefore, beeasily leached.

    At higher p%

    values,conversion to

    E(7%): occurs.

    The resultingnegative charge

    on the molecule

    causes its

    absorption bysoil particles.

    Beaves and

    seeds

    Eoron is reuiredfor upta!e and

    utilisation of 9a*;,membranefunctioning, pollen

    germination, cell

    elongation, celldifferentiation and

    carbohydrate

    translocation.

    Terminal buds are

    damaged, leaving arosette effect on the plant.Beaves are thic!, curled

    and brittle. 2ruits, tubers

    and roots are discolored,crac!ed and flec!ed with

    brown spots.

    9hlorine (9l)

    9hloride (9lF),

    the ionic form of

    chlorine used by

    plants, is usuallyfound in soluble

    forms and is lost

    by leaching..

    verywhere

    "nvolved in osmosis(movement of water

    or solutes in cells),

    the ionic balancenecessary for plants

    to ta!e up mineral

    elements and in

    photosynthesis.

    =educed growth@ stubby

    roots, interveinal

    chlorosis, nonsucculenttissue (in leafy

    vegetables)

    Hunger Signs' Ghen plants do not get sufficient amount of one or more essential nutrient

    elements ,they show poor growth and develop specific deficiency symptoms Ghich is !nown

    as'unger signs.

    Section "C"

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    Ion trans+ortThe plant absorb minerals nutrient from the soil mostly by the rootsbut in

    some plants bylea&es((carni&orous +lants) bromeliads) etc. $ in ionized form.=oots absorb

    some mineral nutrients selectively , including some which may not be essential for them.

    >lants absorb minerals from soil through the root, passes the membrane, in two ways'

    . Passi&e trans+ort'Movement of molecules from high to low concentration. Needs no

    energy input. "ncluding Passi&e di//usion and 8acilitated di//usion

    *. Acti&e trans+ort:3 o&ement o/ molecules /rom lo0 to 'ig' concentration (o++osite

    t'e /lo0 o/ di//usion$. 2eeds in+ut o/ energ* (ATP$.

    >assive transport ' The main theories and concepts of passive transport are '

    . >onann euilibrium t'eor* (>onann ,$' ntry of ions into the cell across theplasma membrane to maintain electrical euilibrium is !nown as ?onann euilibrium

    *. Interce+tion and contact or Ion e1c'ange t'eor*:3 %enn* and 4&erstreet(,-?$ /change of anions and cations absorbed to the root surface with similarly

    charged ions of soil solution is !nown as ion e/change.

    +. ass /lo0 (6ramer ,?$when solute are transported with the conventional flow of

    water from the soil to the plant root under the influence of transpiration .

    ec'anism o/ +assi&e trans+ort

    A. >i//usion'All molecules are in constant motion. As molecules collides into each other,

    directions are changed, causing random dispersal of the molecules. The randommovement of molecules results in diffusion.

    E. >i//usion is t'e mo&ement o/ molecules /rom an area o/ 'ig' concentration to one o/

    lo0 concentration.

    9. This difference in the concentration of molecules across a space is called

    theconcentration gradient)is the difference between the concentration of a solute in

    one place and its concentration in an ad

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    imageH-+

    . ?iffusion occurs due to the

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    in their vacuoles. The solutes cause an influ/ of water. "n this way can plant cells build up

    a large positive internal pressure, the turgor +ressure."t has a decisive influence on the

    maintenance of the rigidity and stability of plant tissues. ach cell e/erts a pressure on itsneighbouring cells. The pressures add up to a large tissue tension.

    imageH-:

    :. H*+ertonic'concentration of solutes outside cell is higher than concentration of solutesinside cell cytosol. Ghen a plant cell is immersed in a'*+ertonic solution,water is

    e/tracted from the protoplasm, it shrin!s. This phenomenon is called +lasmol*sis.The

    process is reversed as soon as the cells are transferred into a hypotonic

    solution (de+lasmol*sis$.

    . Isotonic'. concentrations of solutes outside and inside cell are eual. "n isotonic

    condition)there is no movement of water into the cell of plant. The cell becomes/laccid(lim+$) and the plant may wilt.

    8acilitated di//usion

    >lasma membranes of plant made up of +'os+'oli+id bila*erinterspersed with two type

    of proteins. "t is selectivepermeable and allows the passage of specific molecules needed

    by cell.

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    imageH-*

    Bipid bilayer isim+ermeable to some o/ t'e &ital molecules li

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    "n general, channels are uite s+eci/ic for the type of solute they will transport and

    transport through channels is uite a bit faster than by carrier proteins.

    Three ssential characteristics' . $pecific (selective for single nutrient molecule) *.

    >assive (reuires no input of energy) +. $aturates (nonlinear dependence on

    concentration).

    $ome ion channels are always open, but others have gatesthat open to allow ions to pass

    or close to stop their passage and control t'e c'annel@s +ermeabilit*.The gate can

    controlled by &oltage or ion concentration) e&en b* lig't) 'ormones or ot'er

    stimuli."on channels '

    . Ioltagegated channels

    *. Mechanical gated channels

    +. Bigandgated channels

    imageH-

    Carrier +roteins(also

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    same direction, and anti+orters, where transfer of one solute depends on the

    simultaneous or seuential transfer of a second solute, but in the opposite direction.

    Section ">"

    Acti&e trans+ort (using ATP energ*$B'en a molecule mo&es /rom lo0er concentration

    to 'ig'er concentration ) reuires in+ut o/ energ* (uses ATP$) and is driven active transport systems ' %ydrolysis of AT> ta!es place by

    o Na; ,; transport (sodium pump),Na; is maintained at low concentrations inside

    the cell and ; is at higher concentrations.

    o 9a;; transport

    o %;,; transport iv AE9 transporter superfamily, v osteocast proton pumps.

    *. "on gradient driven active transport systems or9otransport @The gradients of ions(cations or anions) established by AT>ase or by light driven transport may lead to

    secondary active transport of substance li!e amino acid and sugar. These molecules move

    in same directionS*m+ort or in o++osite directions Anti+ort .

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    imageH-H

    "n plants, active transport enables roots to absorb mineral ions from the soil, which are

    therefore more concentrated inside plant cells than in the soil. This reuires AT> energy

    from aerobic respiration, and therefore roots need o/ygen to allow mineral upta!e and a

    waterlogged (thus anaerobic) soil will !ill most roots.

    ;ul< trans+ortBarge molecules, such as polysaccharides and proteins,are too large to

    cross the cell membrane and cross the membrane via vesicles, occurs bye1oc*tosis

    (substances lea&ing t'e cell.$ or endoc*tosis.

    .

    imageH-4

    ineral nutrients u+ta

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    vessels.The passive movement of ions into the apoplast usually occurs through

    ionchannels, the transmembrane proteins that function as selective pores. .

    o S*m+lastic( or inner s+ace $"t is a living continuous system formed by

    cytoplasm and plasmodesmata from epidermis to /ylem parenchyma. An ion enter

    the cell wall of the root hair and pass between the wall and plasma membrane andmoves across the cytoplasm, corte/, endodermis, pericycle through

    plasmodesmata endodermis, a layer of cells that they must pass through to enter

    the /ylem.The entry or e/it of ions to and from the symplast reuires thee/penditure of metabolic energy, which is an process.

    The mineral ions stored in the root hairs pass into /ylem from where these are

    translocated through /ylem along with the ascending stream of water, which is pulled

    up through the plant by transpiration stream. Mineral elements brought to the leaves

    are subseuently assimilated into organic molecules and are redistributed to other

    parts of the plant through the phloem.