LSM3261_Lecture 6--- Growth Responses and Regulation of Growth

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    LSM 3261 Life Form and Function

    Growth Responses and Regulation of Growth

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    Lecture 6 Topics

    Why learn about growth and development?

    Phototropism

    Gravitropism

    Thigmotropism

    Plant hormones and development

    Classical hormones: auxins, gibberellins, cytokinin,ethylene, abscisic acid

    Light signals and plant development

    Photo eriodism Phytochrome

    Responses to herbivores and pathogens

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    Reference

    Solomon, E.P., L.R.Ber and D.W. Martin.

    2011. Biology. 9th ed.

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    Why Learn about Growth and

    Development in this Module?

    Module is about structure and function

    ruc ures n p an s grow an eve op

    through growth responses and regulation of

    grow w c are n uence y e p an sinternal conditions (e.g., genetic makeup) and

    s gna s rom e env ronmen ex erna

    environment) Structures are correlated with functions

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    Tropisms

    Tropism = Directional growth in response to anexternal stimulus such as light, gravity or touch

    Results in an irreversible change in a plant

    part May be positive ornegative, depending of

    whether movement is toward (+) or away (),

    v y, u u Under hormonal control (covered later)

    Types

    Phototropism rav trop sm

    Thigmotropism

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    Phototro ism Directional growth of a plant caused

    li ht

    Most growing shoot tips grow toward

    photosynthesis

    detected by a photoreceptor family What is thecom lementar

    phototropins = Light-activated color of blue?

    -

    enzymes)

    in the blue-light signaling pathway

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    Gravitropism=gravitys direction

    Positive gravitropism means movingtoward the Earths center; negative = inopposite direction

    Root ti s usuall have + ravitro ism

    Shoot tips usually have gravitropism

    Negative (top roots)an pos ve o om

    roots) geotropism in

    tiger orchid

    speciosum)

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    Gravitropism egu a e y =heavy starch-filled granules)

    that settle toward the bottomofspecial cells within the

    root cap in response to

    Amyloplasts reorient asthe root changes direction

    However, recent studies ofArabidopsis thaliana indicate

    a amy op as - ac ngmutant plants still show +

    geotropism More studies needed to

    clarify the mechanism for What can cause a root to changegrav rop sm

    its orientation as it grows throughthe ground?

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    A Corn (Zea mays) Seedling:

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    Thigmotropism Thigmotropism = Growth response to a mechanical

    stimulus such as contact with a solid object

    Twining and curling of stems and tendrils

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    Plant Hormones 1

    Plant hormone = An organic compound that acts as achemical signal eliciting a variety of responses that regulategrowt an eve opment

    Effective at extremely low concentrations (usually

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    Plant Hormones 2

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    How Plant Hormones Act

    Read up model of a signal transduction

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    Auxins

    Auxin = A plant hormone involved in

    development, such as stem elongation,

    ,

    cuttings, e.g., indole acetic acid (IAA) is

    Etymology: Greekauxein, to increase

    CH2COOH

    N

    Indole acetic acid (IAA)

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    Auxins DiscoveryFrancis Darwin

    (1848 1925)

    Charles and Francis= Son of CharlesDarwin; hisassistant

    evidence for this

    n

    Shaded differentCharles Darwin 1809 1882

    coleoptile of canary

    Concluded that aCanary grasssu s ance was passe

    from the tip to the

    ower par s, caus ng

    to bend

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    Coleo tile

    = Protective sheath

    young stem of some

    Protects the plumule

    germination

    o Canary grass

    (Phalariscanariensis)

    o Oat (Avena sativa)Maize seedling

    o Maize (Zea mays)

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    The Darwins and

    Phototropism Opaque covers on the tip

    caused a loss of

    Covers on the lowerportion of the stem had

    no effect on bendingtoward light

    e p was remove ,no bending

    produced in the tip isresponsible for

    p o o rop sm

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    Digression: Research Specialization

    or t ose o you contemp at ng researc as a career Conventional wisdom is to specialize to establish your niche

    op sc en s s researc e var ous areas, e.g.: Charles Darwin: Theory of evolution by natural selection, emotions in

    humans and animals, earthworms, barnacles, power of movement inplants (just covered), orchids, climbers, human evolution, sexualselection, carnivorous plants, etc.

    Albert Einstein: photoelectric effect (Nobel Prize), general theory ofrelativity, relativistic cosmology, capillary action, critical opalescence,unified field theory, etc.

    To scientists chan ed fields e. .: John Maynard Smith (19202004): aeronautical engineer then took

    second degree in genetics, becoming one of the most famous, .

    More current thinking: Become a deep generalist

    Next best thing: collaborate, but must be able to be a team player,have social skills

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    Avena (Oat) and Auxin

    coleoptiles in the 1920s Placed coleoptiles on agar blocks

    Phototropic = light-responding

    ,

    Agar blocks caused increased growth on the side of the growing shoot

    Frits Warmolt

    Went (19031990)

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    Auxin Effects

    The movement of auxin is polar,(unidirectional, downward from its site

    of production, usually shoot apicalmeristem) resulting in an auxin gradientthat tri ers s ecific res onses

    Effects include:

    oApical dominance

    o ee ess ru s

    oOthers

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    Cell Elongation

    elongation in stemsand coleoptiles

    Movementof auxin

    Auxins cause theincrease in cell length

    on the side awa fromlight

    Causes bending

    Indole acetic acid(IAA) is one of themost im ortantauxins

    Causes acidification,an so en ng, an soelongation withturgor pressure, of thecell wall

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    Apical Dominance =

    The inhibition ofaxillary bud growth by

    Auxin produced in

    inhibits axillary budsnear the apex from Mango tree

    Leading branch then

    ( Mangifera indica)

    the rest of the plant

    Scots pine tree

    (Pinus sylvestris)

    Ethylene and cytokinin Do shrubs have

    dominance

    strong or wea ap ca

    dominance?

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    Seedless Fruits

    flowers of certain

    ,

    fertilization leads to

    ,

    e.g.,Parthenocar ic = Fruit roduction without fertilization

    Not all seedless fruits,

    e.g.,oThomson seedless

    grapes (abortion of

    embryos after

    fertilization, application

    of gibberellin)

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    Synthetic Auxins

    is a naturalproduct

    Synthetic = humanmanufactured, e.g.,

    Naphthalene aceticac or rooinduction

    -

    dichlorophenoxyacetic acid (2, 4-D) =eu co er c e

    2, 4, 5-

    acetic acid (2, 4, 5-T)= eudicot herbicide

    The US Army sprayed Agent Orange (2, 4-D + 2, 4, 5-T +

    Others picloram [systemic herbicide]) on Vietnamese forests todefoliate them (toxic dioxin was a contaminant): possible linkto serious illnesses in Vietnam war veterans and Vietnamese

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    Digression: Use of Scientific Research

    Scientific knowledge is neither good nor bad

    Its use is determined by humans

    For ood e. . a riculture to im rove ieldspropagate plants, cure human diseases, nuclearpower for electricity generation

    For bad, e.g., defoliate forests for war, biologicalwarfare, weapons of mass destruction

    Always important to behave ethically and

    morall in the a lication of science However, not always easy to decide what is

    ,

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    Tutorial Assignment 3 and Practical 3

    T urs ay, 29 ep 2011 Go first to LS Lab 7

    o Tutorial Assignment 3 briefing: 2.00 to 2.45 pm

    e r p o ngapore o an c ar ens:

    3.00 to 5.30 pm

    oDress comfortably, bring umbrella/rain coat,water, notebook, schedule, camera, insect

    repellent

    o Leavin from Science Drive 4 behind UniversitHall at 3.10 pm

    .

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    Gibberellins GAs Eiichi Kurosawa (Japan) in 1926

    studied a disease of rice (Oryzasativa that caused exa eratedgrowth caused by a funguscalled Gibberella fujikuroi

    and UK showed giberellins areproduced by healthy plants

    can be extraordinary Causebolting rapid growth of

    -plants, prominent in cabbage-related species (Brassicaceae)

    e s are s mu a e o e onga eAND divide

    Stimulate flowering andRice mutant which

    germ na on n many p an s

    Can bypass requirement forvernalization in many plants

    gibberellinNormal rice

    li i

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    Vernalization=

    exposure to low temperatures,

    e.g., germination or flowering

    Adaptation to prevent

    germination or flowering in

    GAs can substitute for the low

    tem erature treatment

    Can induce flowering inbiennials in first year, e.g., carrot

    or perennials, e.g., plum blossom

    (Prunus mume) needs 550800

    hours of chillin Can induce seed germination in

    many temperate species, e.g.,

    apple

    B l i

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    Bolting

    In the first yearIndian blanket

    (Gaillardia

    pulchella) grows as arose e o eaves

    from a shortened

    In the second year ofrowth boltin

    occurs (right) with

    stem elongation

    Is the result of

    gibberellin

    pro uc on

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    CytokininsCytokinins promote

    = Plant hormone involved in variousaspects of plant growth and

    Normal tobacco plant

    cytokinesis (cell division)

    ,

    of senescence, and differentiation

    plant tissues in culture

    cytokinin and auxin High ratio of cytokinin to auxin promotes

    shoot formation

    Low ratio of cytokinin to auxin promotes

    In apical dominance, auxin inhibits axillary

    bud formation; cytokinin promotes it

    Cytokinins delay cellular aging, or

    senescence High cytokinin transgenictobacco plant

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    and Auxin

    Control of

    activityby auxin

    and c tokinin

    Determination of thedifferentiated tissue

    type depends on the

    ratio of auxin to

    cy o n n

    h l 1

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    Ethylene 1

    Ethylene gas (C2H4) isproduced by plants

    = Gaseous plant hormone

    involved in various aspects ofplant growth and development

    such as leaf abscission and

    fruit ripening Is critical to many plant

    functionso Inhibits cell elongation

    o Promotes see germ nat on

    o Promotes apical dominance

    o Fruit ripening

    4

    wrappers are needed for keeping fruitsfrom ripening in transit

    E h l 2

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    Ethylene 2

    Ripening fruit produceethylene, so a ripe fruit maypromo e r pen ng o near yfruit

    green then ripened with ethylene

    o Ripe banana in a bag with otherfruit ma romote ri enin of the

    second fruito One rotten apple spoils the wholebarrel: because of ethylene

    Leaf abscission indeciduous plant promoted ast e et y ene : aux n rat o ncells at the base of the

    fall (autumn)

    E h l 3

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    Ethylene 3

    Thigmomorphogenesis =Plant developmental

    responses to mechanical

    stimuli, e.g., wind, rain,contact with passing animals

    Mechanically disturbed

    plants produce additionalethylene whicho Inhibits cell elongation

    o Enhances cell wall thickeningJuniper trees

    a es s ems s or er an

    thicker so more resistance

    o amage

    Ab i i A id (ABA)

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    Abscisic Acid (ABA)

    Abscisic acid is present in large amounts in dormant seeds;

    concentration drops as it is washed out

    Gibberellin or cytokinins used to overcome dormancy

    Promotes plant changes during water stress

    Causes stomata to close, saving water

    In a abscisic acid-free maize mutant, a

    seed germinates while still on the plant

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    Ph t i di d Fl i

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    Photoperiodism and Flowering

    Photoperiodism = Anyresponse of a plant to the

    re at ve engt s o ay g t

    and darkness in the duration

    Plants types

    =- -

    Long-day (= short-night)

    -

    Day-neutral

    Short Day PlantsChrysanthemum

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    Short-Day Plantsy

    Flowerwhen night is longerthan a critical period (long-n g t p ant)

    Typically flower in late summero nsett a

    Cocklebur

    Long Day Plants

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    Long-Day Plants

    Flower when night ish r r h n ri i l

    period (short-night plant)

    Flower in late s rin or

    Black-eyed Susan

    early summer

    thaliana

    p nac

    Types of Short and Long Day Plants

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    Types of Short- and Long-Day Plants

    Types

    period of daylight (all or none effect depending on duration

    of daylight per day)o Quantitative or facultative: flowering accelerated by

    specific period of daylight per day

    o o easy o scr m na e e ween e wo ypes ecause a

    photoperiodic requirement may be more pronounced under, . .,

    photoperiodic requirement under one temperature and have a

    facultative photoperiodic requirement under anothertemperature

    o Better to treat these two categories as two ends of a

    con nuum

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    -

    Long-Day

    Plants The behavior can

    be modified by a

    flash of red light

    = long-

    night

    This indicates theim ortance of a

    p ant

    red-sensitive

    pigment system =short-

    night

    plant

    I t di t D Pl t

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    Intermediate-Day Plants

    Flower when exposed to an

    Sugar cane

    o eus

    ,

    typical sugar cane plant to flower in the town of Pontianak(Indonesia) (situated exactly at the equator)?

    Day-Neutral Plants

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    Day Neutral Plants

    Flowering not initiated in

    response o seasona c anges

    in day- and night-length

    ower n response o o er

    external or internal stimuli

    Not All Hormonal Responses

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    Not All Hormonal Responses

    re u y arac er ze Long-day tobaccop an cot anasylvestris) is grafted

    onto a day-neutraltobacco plant(Nicotiana tabacum)will induce lon -da

    behavior, causingboth plants toflower whenexposed to long days

    An unknown

    ,is thought to beresponsible for theresponse

    Phytochrome Detects

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    y

    The photoreceptor pigment for

    photoperiod is called phytochrome =

    Family of about 5 blue-green pigment

    proteins (each with own gene)

    structures not

    examinable!

    complementary color of red?

    There are 5 kinds of these pigments

    in all vascular plants studied so far

    phyA, phyB, phyC, phyD and

    h E in rabido sis thaliana

    phyB important at all stages;

    others important over narrow

    ranges o e e cyc e

    phyA antagonizes phyB: in long-

    day plants, phyB inhibits

    flowering while phyA may induce

    flowering

    Electromagnetic Spectrum

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    Electromagnetic Spectrum e -a sor ng nm p y oc rome r a erna es w ar-

    red-absorbing (730 nm) phytochrome (Pfr)

    Phytochrome Shifts between 2 Forms

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    Phytochrome Shifts between 2 Forms

    Red light (660 nm) converts Pr to Pfr, and far-red light (730 nm)converts Pfr to Pr

    Conversion of Pr to Pfr triggers or inhibits physiological responseslike flowering

    Phytochrome Effects

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    Phytochrome Effects

    Shade avoidance

    Germination

    eep movemen s n eaves

    Shoot dormancy

    Leaf abscission ,

    fruits, and leaves

    Shade Avoidance 1

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    Shade Avoidance 1

    = Response in which plants tend togrow taller when shaded to compete

    Neighboring plants absorb more

    Lamina is like a light filter

    Shade Avoidance 2

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    Increasing the ratio of far-red to red light increases

    In tomatoes, increased far-

    fruit production

    Germination

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    Some species have seeds with a light requirement

    involving phytochrome

    M x li h i h hi h r d f r-r dratio to germinate

    Examples

    o emperate ec uous orest n spr ng n as tt e s a e,

    hence good time for such seeds to germinate

    o oneer ree spec es n e rop cs germ na e on y un eropen conditions where such species thrive

    Circadian Rhythms

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    y Nearly all organisms display an internal clock

    Approximates 24-hour cycle

    , ,

    etc.

    In plants circadian rhythms common ause open ng c os ng o s oma a Movements of leaves called sleep movements

    Rain tree (=pukul lima)

    (Samanea saman)

    ean aseo us

    vulgaris) plant

    During the day (left)

    At night (right)

    Settin the Biolo ical Clock

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    Settin the Biolo ical Clock

    - -

    implicated in setting the biological clock for many plants

    homologous with those of clock proteins in fruit flies,

    ,

    the fruit fly and mouse

    o og ca c oc s are very mpor an n coup ng e

    activity of plants to important events, e.g.,

    o Flower opening must be coordinated with pollinators

    Plant Responses to Herbivores and Pathogens

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    Plant Responses to Herbivores and Pathogens

    Plants have an innate immune

    system to fight local infections, but

    this immune system is unlike thosein animals

    A signal transduction pathway is

    tr ggere w en a mo ecu e rom a

    pathogen or herbivore binds to a

    A hypersensitive response results

    and this seals off the infected areacausing it to die to create a necrotic

    lesion (dead area) to prevent or

    slow the spread of the infection

    S stemic Ac uired Resistance SAR

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    S stemic Ac uired Resistance SAR

    Plant also sends signals from the site of

    oca ze n ec on roug e p oem

    These lead to SAR = Buildup of long-

    lasting enhanced resistance throughout

    the plant

    SAR is broad spectrum, heighteneddefense against many pathogen types

    and not only the causal pathogen

    Such hormone-si nallin moleculesinclude:

    o Jasmonic acid

    o

    Methyl salicylate (oil of wintergreen)