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Hormones
Chemical signals that control plant growth and development
Diverse group of chemicals that have the ability to control the growth and developmental processes in plants (and animals)
They can be modified amino acids, small organic acids, terpenoids, gases, steroids etc.. They are present in very low concentrations in cells, but control growth and development of plants.
Auxin
Controls many physiological processes of plants
-Shoots always grow towards light
-Roots always grow towards gravity
-Controls proper fruit growth of strawberries
-apical dominance. Shoot apical meristem dominates over axillary buds due to high level of auxin
Major auxin found in plants
Indole-3 acetic acid (IAA)
Auxin synthesis
Synthesized inyoung tissues ike SAM. Then, auxin transports toward roots generating a gradient of auxin from SAM to root (this is known as polar auxin transport).
Morphogen
This gradient of auxin is necessary to maintain the architecture (shape) of the plant. Thus, auxin also considered as a "morphogen".
Polar transport of auxin
Due to two different types of transporter proteins called influx and efflux carriers
Auxin is transported by membrane transporter proteins
Influx carrier
Efflux carrier
Influx carrier
Transports auxin into the cell
Efflux carrier
Transports auxin out of the cell
Auxin is ____ during tropic responses
Redistributed
Phototropic response and auxin
Light changes direction of auxin movement
Cells with more auxin elongate more causing bending of the stem
Light changes the localization ofEfflux carrier (transporter) changing the direction of auxin transport. Blue light is required for this response.
Gravitropic response
Auxin also involve in gravitropic response.
More auxin is transported to cells in lower side causing cells in the upper side of the root to expand.
This is kind of opposite response compared to what is happening in the shoot (phototropism).
Auxin concentration is higher in what region of the root
Columella
Statoliths
Large starch granules that rerceives gravity in roots
Columella cells contain bigger amyloplasts (starch granules) They are also called statoliths.
Statoliths rest on ER membranes of the lower side of the cell due to gravity.
Starch-statolith hypothesis of gravity sensing
Changing the direction of gravity leads to movement of statoliths to a new location (lower ER membrane). This generates a mechanical pressure on ER membrane.
How does auxin control plant growth and development
Genomic responses and non genomic responses
Genomic response for auxin control
Generally, auxin controls the expression of many genes that produces proteins necessary to control growth and development.
Non genomic responses of auxin control
Also, auxin controls the activity of some proteins through post translational modifications to control growth and development
How does auxin control gene expression
High increase of auxin promotes the degradation of the repressor protein.
Degradation of the repressor protein allows the proper interaction of TF leading to gene transcription
Repressors are degraded through Ubiquitin-Proteasome Pathway at high concentration of auxin
Agricultural uses of auxin
Chemicals with auxin activity are being used as selective herbicides.
Auxin is used in tissue culture as it can control cell division, expansion and cell differentiation.
Auxin is also used to produce parthenocarpic (seedless) fruits.
Rooting powder mainly contains auxin
Natural use of auxin as a herbicide?
Cant be used because plants have the means for degrading or metabolizing IAA.
Synthetic auxins cannot be degraded by plants as there are no natural mechanisms to metabolize them. thus when used at higher concentrations they kill dicot plants
Gibberellic acid (GA)
Many physiological responses are regulated by GA
GA controls thestem elongationof plants. Mutants that do not produce enough GA show short stems
GA induces stem elongation of rosette plants
GA is used to enhance the growth of seedless grape
GA induces early reproductive growth of plants
GA promoting seed germination
By controlling the expression of genes that are required for seed germination
Germinating seeds and energy
When seeds germinate until they produces leaves for photosynthesis, germinating seedlings have to rely on glucose produced in the seed. Carbohydrates are stored in seeds as starch (a complex carbohydrate). Starch should be hydrolyzed to glucose to be used as an energy source as well as to produce other essential macromolecules.
How does GA control alpha-amylase gene expression
GA first induces the expression of myb gene to produce myb protein.
Myb protein is a transcriptional factor that interacts with enhancers ofα-amylase gene.
DELLA repressors and high levels of GA
DELLA repressors are degraded through Ubiquitin-Proteasome Pathway at high levels of GA
Developmental processes that ethylene regulates
Fruit ripening
Senescence
Induces root hair formation
In dark, ethylene inhibits hypocotyl elongation, thick hypocotyl and exaggerates apical hook
How does ethylene control developmental processes
Ethylene is a gaseous hormone.
Like other two hormones, ethylene also control the expression of specific set of genes that are required for ethylene induced developmental processes.
Ethylene also uses ubiquitin-proteasome pathway to control gene expression.
However, unlike auxin and GA (in which repressor proteins are degraded), ethylene inhibits the degradation of a transcriptional factors (TFs), which are degraded in the absence of ethylene.
Ethylene activates gene expression by inhibiting the degradation of TFs
When there is no ethylene, TFs necessary for the induction of genes (by ethylene) are continuously degraded. So genes cannot be activated.
When there is ethylene, TFs necessary for the induction of genes (by ethylene) are accumulated as ethylene will inhibits the degradation of TFs.
Gene induction depends on the abundance of the
How TFs are degraded through Ubiquitin-Proteasome Pathways
When there is no ethylene signaling pathway is active and TFs are degraded through ubiquitin Proteasome pathway.
When there is ethylene, ethylene binds with its receptor located in the ER membrane and inhibits the signaling pathway, thus inhibiting the degradation of TFs. TFs now can induce ethylene responsive genes.
Gene induction depends on the
Abundance/activity of the TF or Repressors
How auxin and GA activate gene expression
By degrading repressor proteins
Two major steps of E3 (SCF) mediated protein degradation
1) Ubiquitination of target proteins
2) Degradation of the target proteins
5 steps of E3 mediated protein degradation
1. Ubiquitin undergoes ATP-dependent activation by E1
2. Ubiquitin is transferred to E2
3. Ubiquitinated E2 forms a complex with E3 ligase and the target protein
4. The target protein is ubiquitinated by the E2-E3 complex
5. The target protein is degraded by the 26S proteasome
Specificity of the pathway is regulated by
E3 (SCF) complex
Auxin summary
Molecular glue between Aux/IAA & F-box protein
GA summary
Conformational change in DELLA protein?
Ethylene
Conformational change in EIN3 protein?
LECTURE 7
Three phases of plant life cycle
Embryogenesis
Vegetative phase
Reproductive phase
Embryogenesis
Development of the embryo from the zygote
Vegetative phase
Development of a mature plant from the embryo
Reproductive phase
Development of reproductive structures
Two generations that plant life cycle alternates between
Sporophyte 2n
Gametophyte n
Development of the sporophyte
In flowering plants, sporophyte is easily visible
When a seed germinates, it produces the sporophyte
Sporophyte produces the reproductive structure which is the flower
Process:
Fertilization
Zygote develops into embryo 2n?
Ovary develops into fruit, ovule develops into seed
Germination
Young sporophyte
Flowers with pistil and/or stamen
Pistil has ovary, ovule, megastore mother cell undergoes meiosis
Stamen has microspore mother cells undergoing meiosis
One of the 2n cells in the ovary will become
Megastore mother cell
Cells in the tapetum of the anther will become
Microspore mother cells
The male gametophytes of flowering plants are also referred to as _____.
pollen grains
Development of gametophyte
Microspore mother cell divides by meiosis to produce 4 equal haploid cells, called Microspore. Each microspore divides by mitosis to produce two cells- pollen (male gametophytes)
Development of megaspore
Megaspore mother cell divides by meiosis to produce 4 equal haploid cells, megaspores. 3 of these will degenerate leaving only one megaspore
Meagaposre divides by mitosis to produce 8 nuclei that will end up generating 7 cells that comprise the female gametophyte
Pollination
When pollen comes into contact with stigma of the pistil
Pollen tube
Pollen germinates via pollination and produces a pollen tube, which is guided by tube nucleu to the ovary (towards the egg)
Generative nucleus divides by mitosis to produce two what
Sperm nuclei
One sperm nucleus fuses with egg cell to produce___. Other spern cell fuses with ____ to produce 3n cell
2n zygote
Polar nuclei
Zygote will divide by mitosis to produce ____. 3n cell divides by mitosis to produce ___
Embryo
Endosperm
Establishment of polarity
Zygote will divide by mitosis to produce embryo. 1st cell division is a centric to produce an applicable smaller cell and longer basal cell. This is because of polarity establishment in the zygote probably through auxin distribution
Several distinct development stages that embryogenesis goes through
Embryo sac
Zygote
2-cell
8-cell
16-cell
Early globular
Late globular
Transition
Late heart
Seedling
Apical cell divides to produce ___ while basal cell divides to produce ____ and ____
Embryo proper (globular structure)
Uppermost cell (hypophysis)
Lower suspensor
Hypophysis produces ___ while suspensor ____
Produces the root
Helps absorb nutrients from the endosperm to the growing embryo
Role of auxin in embryo development
During the different stages of embryo development, we can observe maximum auxin concentration in different tissues where we can see higher cell division
What inhibits auxin biosynthesis
Trans-cinnamic acid
Whawt it's he auxin effluent carrier
Pin1
Mutations in auxin related genes cause
Defects in embryo development
How mutations in auxin related genes cause defects in embryo development
When the GNOM gene is mutated, embryo files to develop both shoots and roots
When MONOPTEROUS gene is mutated, embryo families to develop roots
MONOPTEROUS gene encodes a repressor protein involved in auxin response
Apical-basal axis growth is due to
SAM and RAM
Radial axis growth is due to
cambium and cork cambium
Growth in radial axis is due to what cell divisions
Periclinal and anticlinal
Periclinal division
Represents cell division parallel to the periphery of teh stem
Anticlinal division
represents cell division right angle to the periphery of the stem
Majority of plant growth and development occurs through
Post embryonic development
LECTURE 8
Seeds are produced in the ___. After fertilization, ___ turns into the seed while the ___ develops into a fruit
Ovary
Ovule
Ovary
Most of the space of the embryo sac will develop into the ____ while the fertilized egg (zygote) develops into the ___
Endosperm
Embryo
In some dicots, content of the endosperm will do what?
Be absorbed into the growing cotyledon, which replaces the endosperm
Seeds are an adaptation by plants to
The terrestrial environment
Testa
Seed coat
Aleurone layer
Seeds with endosperms contain this
Produces GA
Endospermic
Plant that contains entire endosperm
Non-endospermic
Endosperm absorbed by embryo
Dormancy
If a live seed does not undergo germination when exposed to suitable environmental conditions, it is considered in dormancy
Primary dormancy
Seeds do not germinate even in favorable environment during the dormancy period
Secondary dormancy
Even after dormancy period seeds may not germinate if unfavorable environment conditions are present
Primary dormancy could be due to
Exogenous dormancy
Endogenous dormancy
Exogenous dormancy
Seed coat-imposed dormancy (limits water, gas exchange, inhibitors, mechanical constrain)
Endogenous dormancy
This is due to the germination inhibitory hormone, ABA produced by the embryo
Viviparous
Seeds do not have a dormancy period and germinate on the mother plant
An adaptation to environmental conditions
Can viviparous germination be seen on non-viviparous plants?
Yes, this is due to mutations of genes that are required for dormancy
What hormones regulate dormancy/germiantion of seeds
ABA
GA
ABA promotes
Dormancy
Biosynthesis of ABA
During seed maturation process, water content will drastically reduce. This will enhance ABA biosynthesis. In addition, sensitivity to ABVA can also be increased
GA promotes
Seed germination
Biosynthesis of GA
During seed imbibition cells absorb water, this will enhance GA biosynthesis. In addition, sensitivity to GA can also be increased
Breaking dormancy different requirements
Light
Chilling
After-ripening period
Some chemicals (by inhibiting ABA response)
Mobilization of food storage during seed germination
During seed germination, complex storage food such as starch is converted to simple sugars (glucose) for the initial growth and development
Starch to glucose conversion requires a-amylase enzyme that is produced in the aleurone layer
GA induced starch hydrolysis
Occurs through a-amylase induction in aleurone layer. GA first induces MYB gene and MYB protein functions as a TF to enhance a-amylase gene
Epigeal
pertaining to growing on or germinating above the surface of the earth (depends on the position of cotyledon after germination)
Hypogeal
Growing/germinating below the surface of the ground (depends on position of cotyledons after germination)
What promotes stem growth during seedling growth
Auxin
Acid growth hypothesis
Auxin promotes H+ secretion from cells leading to acidification of cell walls. This will help to increase enzyme activity to degrade cell walls, which leads to cell wall loosening due to turf or pressure causing cell expansion.
LECTURE 9
Reproductive development is regulated by what two major factors
Endogenous
Exogenous