Signaling Systems Within The Plant: A New Era of …...Plant Signaling Systems •Plants are good at...
Transcript of Signaling Systems Within The Plant: A New Era of …...Plant Signaling Systems •Plants are good at...
Signaling Systems Within The
Plant: A New Era of Under-
standing
Phil Thomas
Senior Agri-Coach
Agri-Trend Agrology
Factors affecting Yield and
Quality
Yield
• Soil Type
• Organic Matter
• Soil Texture
• Soil pH
• Soil Aggregate Size
• Rooting Depth
• Field Topography
• Ion & Cation Exchange
Capacity
• N, P, K, S, Ca, B, Cl, Mg, Mn,
Mo, Fe, Zn levels
• Field location /latitude
• Soil Water Holding Capacity
• Soil Aeration
• Soil Water Infiltration Rates
• Soil Crusting
• Soil Drainage
• Field Slope
• Rain & Snow
• Solar Radiation
• Evapotranspiration
• Carbon Dioxide
• Relative Humidity
• Wind Velocity
• Soil & Air Temperature
• Frost Free Days
• Oxygen
• Hail
• Flooding
• Rotation – Crop &
Pesticide
• Seedbed Firmness
• Pests – Weed, Insects &
Diseases
• Harvest/Swath Date
• Variety
• Seed Quality
• Seed Treatment
• Seeding Date, Rate, Depth
& Speed
• Pesticide – Choice &
Timing
• Tillage Intensity
• Fertilizer – Source, Rate,
Placement & Timing
Management of Factors • Due to multiple “Stress” factors
crops do not reach their genetic
yield potential
• Plants are continually exposed to
abiotic and biotic stresses that
negatively influence growth and
development
What is the most important input to
crop production?
Maximizing Photosynthetic Efficiency
2CO2 + 2H2O → C6H12O6+ 6O2
Nitrogen
Calcium
Potassium
Sulfur
Nitrogen
Phosphorus
Magnesium
Oxygen Carbon
Oxygen
Zinc
Copper
Boron
Manganese
Iron
Hydrogen
Chloride
Molybdenum
Life Cycle in Plant Cells
• Most plant life cycle
processes occur at
the plant cell level
regulated by genes.
• Every biochemical
activity in a plant is
regulated by genes!
1. Photosynthesis Sugars
Carbohydrates
(including starch)
2. Respiration (Krebs Cycle)
Makes
ATP Energy
Amino Acids
CO2
H2O
CO2 & H2O O2
+ N
Cellulose Lignins Proteins
+ S
•Nucleic Acids
- DNA, RNA
•Chlorophyll
•Growth Regulators
•Etc.
4. Transpiration
3.S
yn
thesis
Lipids
How a plant works at the molecular level
5.T
ran
s -
locati
on
Think like
a plant
Plant Genetics, Physiology and
Biochemistry
• What’s really changed in the past 10
years is:
• the shift from Genomics (number and
types of genes involved)
• to Proteomics (protein products of the
genes)
• and now Metabolomics (metabolites).
Genes Regulate
Vegetative growth:
Germination,
seedlings,
shoots, roots and
branches
Reproductive growth:
# of buds, flowers,
pods, seeds, and
seeds per pod
Yield:
Biomass, plant height, dry
matter, drought stress, heat
stress, and leaf area index
Genes Regulate
Chemical
composition:
Hormones:
Abscisic acid
Auxins
Cytokinins
Ethylene
Gibberellins
Others
For lignin content,
Oil/protein content,
Carbohydrates
And amino acids
Proline content:
(mineral elements)
Cytoplasmic streaming
Improved
enzyme
activity
Genes Regulate
Photosythesis
efficiency:
Leaf assimilation
Total chlorophyll
content and
intensity
Water use efficiency
Water management -
through lower stomata resistance
Higher intensity of transpiration
Increased water uptake by roots
Plant signaling Systems (genes)
Hormones
Peptides Or
PGR’s
Macro-proteins which are snippets of mRNA sent from cell to cell via the phloem. (transport and defence against various stresses)
Plant signaling genes
Hormones Peptides Or
PGR’s
Signals are
secreted in
response to
environmental
factors (nutrient
abundance,
drought, light,
temperature,
chemical or
physical stress)
Germination, Rooting, Growth, Flowering, Foliage, Death
Ho
rmo
ne L
evels
Key Nutrient
Hormone
Co-factors:
GA
STAGE II:
Vegetative
Growth
B, Ca, Cu, Fe, K, Mg,
Mn, Zn, amine N
Cell
Sizing
Cytokinin
STAGE I:
Germination &
Establishment
Ca, Fe, Mg, Mn, N,
P, Zn, B
Cell Initiation[ Cell Division ]
Ethylene
STAGE III:
Flowering &
Reproduction
B, Ca, Cu, K, Mg, Mo,
amine N
Cell
Maturity
ABA
STAGE IV:
Fruit Sizing &
Maturity
B, Cu, K, Mg, Mn, Mo, P
amine N
Senescence
Auxin
Seed Germination Seeds imbibe 45% of their weight in
water which leads to swelling and
seed coat breakage.
Genes activated hydrolytic
enzymes break down stored oils and
proteins into chemicals (+oxygen)
for metabolism and growth.
Oxygen is used in aerobic
respiration for energy until the plant
has leaves. Once the radicle
emerges germination ends.
Cytokinin (CK) • Root tips = factories of cytokinin
synthesis (involved in hormone
metabolism, up-take of macro-nutrients,
protein synthesis + morphological
response)
• Nitrate, sulfate and phosphate stimulates
CK production in the roots and later in
the shoots. In leaves CK involved in
stomata opening + the above root factors.
Plant Signaling Systems
• Plants are good at math!
• Receptors (internal clock) within the leaves at
night calculate amounts of starch available and
estimate time to dawn.
• Signals adjust rate of consumption so the leaves
don’t starve from lack of energy until the sun
returns.
• This helps the plants continue to grow in the
dark
Britain’s John Innes Centre – Journal of eLife
Plant Signaling Systems
• Signaling systems allow plants to see,
smell and feel (also movement).
• Plants have the ability to sense the
environment and adjust their
morphology, physiology and phenotype
accordingly.
Plant Signaling Systems
• Plants perceive and can react to stimuli
such as chemicals, gravity, light,
moisture, infections, temperature, oxygen
and carbon dioxide concentrations,
parasite infestation, physical disruption,
and touch. Plants have a variety of means
to detect such stimuli and a variety of
reaction responses.
Plant Signaling Systems (smell)
• For example a willow tree branch being
attacked by tent caterpillars produces
salicylic acid (SA) which makes its leaves
taste bitter and unpalatable. This signal
of SA is also released into the air and
detected (smelled) by nearby branches or
trees which then also produce SA
providing protection.
• Lima beans attacked by an insect or
bacteria do it too.
Plant Signaling Systems (smell)
• Wounded tomatoes are known to
produce the volatile odour methyl-
jasmonate as an alarm-signal when
attacked. Plants in the neighbourhood
can then detect the chemical and prepare
for the attack by producing this or other
chemicals that defend against insects or
attract predators.[5]
Plant Signaling Systems (feel)
• The effect of
touching a Mimosa
plant with fingers
causing the leaves
to rapidly fold up.
• The cocklebur weed
can die simply by
touching it a few
seconds for a few
days
Photo Receptors
• These light sensors detect shading from
neighbours and produce a signal “auxin”
which causes some plants to grow taller.
• There are likely over 300 kinase proteins
in canola with many diverse functions
and pathways. These signals are involved
in developmental and defence functions.
(P-NB)
Plant Signaling Systems
(movement)
• For example
sunflowers with
photo receptors
send signals to
slowly move heads
with the sun.
• Another is the
Venus fly trap.
Canola Life Cycle
Plant Signaling Systems • Plants can see UV
light (red & blue)
via photo receptor
proteins.
• Canola plants need
over 10 to 12 hours
of day length/day
before the
reproductive stage
processes start.
GDD’s (0 base)
Reproduction – B is essential
Hours after plant receptors find required DL + GDD’s
FT gene in all leaves makes a
signal molecule called FT
protein.
Transported by phloem to growing tip
combines with and
activates a FD
protein from an FD
gene.
FT/FD signal
Acts on genes - turns stem cells into flower buds
Canola Sex - Flowering
Immature Bud
Immature Stigma
Immature anthers
Canola Sex - Flowering
Flower opens
within 2-3 days pollen
is produced by
anthers
Canola Sex - Flowering
Anthers mature and release pollen
Stigma tip has adhesive to capture
pollen
Canola Sex - Flowering
Pollen land on the
stigma and absorbs
water and nutrients
from the stigma to
germinate and form a
pollen tube
Over 100 pollen grains are required on the stigma to
fertilize all the ovules
GABA Key signalling molecule that triggers pollination – Boron is a precursor
Stigma
Ovules
1st ovule that matures sends out the GABA signal
once 1st ovule is pollinated
the next ovary starts to
send the signal etc
Pollen tube follows the signal and pollenates the 1st ovule
Pollen grain
Takes only a couple of days or less for each flower for complete pollination
Pollen Tube
If boron is deficient the tip of the pollen
tubes will burst and no pollination will occur
Flowering
• From the start to the end of flowering
there are about 8 to 9 Gibberellins
involved that have effects on seed
formation, proteins and oil.
• Gibberellins are also critical for root
growth as they regulate numbers of cells
and their size.
• K and Zn essential
Plants on Steroids • Brassinosteroids from chloroplasts, join a
protein on the surface of a plant cell and
send signals to the cell's nucleus causing
plant genes to be expressed. (P is
important as the signal is a protein
phosphatase).
• Chemical signals activate a cascade of
gene activity regulating growth and
development (response to gravity, light,
& resist stresses).
Plant Signaling Systems • Water movement is controlled by a cell
membrane gene that produces an
aquaporin (macro-protein) signal. This
signal opens or closes cell membrane
water/protein channels when drought or
waterlogging occurs resulting in
improved water use efficiency and
movement.
• There are about 35 aquaporin’s in plants.
• Phosphorus is important
Water Management
• Understanding how roots grow and how
hormones control that growth is crucial
to improving crop yields.
• A gibberellin protein signal plays a
crucial role in controlling the size of the
root meristem, and that it is the
endodermis which sets the pace for
expansion rates in the other root tissues.
Water Management
• Carbon is essential
for all life!
• CO2 critical for
photosynthesis.
• Plants open stomata
during the day – a
problem - take in
CO2 but lose water
vapour.
Water Management
• When roots sense a
water shortage they
send a macro-
protein signal to
produce absicsic
acid (ABA) that’s
translocated to the
leaves which closes
the stomata.
• K & Ca are NB
Water Management • ABA triggers a signalling cascade in
stomatal guard cells which closes the
stomata reducing water loss.
• ABA induces the production of H2O2 in
guard cells which then activate calcium
channels – Both ABA and H2O2 induced
Ca channels are important mechanisms
for stomata closing.
• Therefore K and Ca are important
Signals Involved with Nutrients
• It is important to understand the
molecular basis of nutrient uptake and
transport within the plant and the genes
responsible
• Uptake and transport of nutrients are
gene regulated by signals
Signals Involved with Nutrients
• For example there are 14 genes (that we
know of) involved in the transporters
(uptake and transport) of sulphur within
the plant
• These genes are in 5 groups (in the roots,
leaves, stems and cell to cell)
• There are many more genes regulating
transporters of Mg (7) N (37) and P (142)
Signals Involved with Nutrients
• Where S is deficient = big yield loss
• But if S is added then uptake of Se and
Mo is decreased due to the sulfate
transporter expression and competition
from Mg, N & P transporter genes.
• But if Mg, N & P up-regulated genes are
switched on S uptake is increased
Nutrients Involved with Signals
• Nutrients are critical for the production
of signals and/or for the products
produced (metabolites, enzymes, etc.)
• For example biological nutrients are key
for making enzymes:
• Ni (2) Mo (4) Mn (50)
• Fe (100) Cu (100) Zn (1200)
Nutrients Involved with Signals • While Mo is only involved with 4
enzymes it is critical for plant growth
– Nitrate reductase – no N nutrition without
Mo
– Sulfite oxidase – NB in chloroplasts
– Zanthize dehydrogenase – disease defense
– Aldehyde oxidase – NB in hormone signals
metabolism (cytokinins, ABA, Auxins, ROS
• Mo is a cofactor for Moco sulfurase (MG,
S & Fe) a protein for drought tolerance.
Nutrients Involved with Signals
• There are a large # of genes that regulate
molecular processes that respond to K.
• Jasmonic acid – a defence metabolite
• K is involved in signals for accumulation
of secondary Jasmonic defense
metabolites for fungal, bacterial, viral
and insects attacks. Therefore, plants are
healthier.
Nutrients Involved with Signals
• Cu, Zn & Mn are co-factors for a large
number of enzymes such as SA which
results in “systemic acquired resistance”
improving disease resistance.
• There is a synergism with seed
treatments and seed quality, health and
yield.
Protection Response
• Plant with no stress (weather, nutrients,
disease, water, good rooting depth, etc)
may only have 3 genes (turned on or off)
so there is limited resistance activity.
• But in plants with stress there may be 60
to 70 genes which are strongly affected
under stress (and turned on).
Protection Response
• SAR – “systemic acquired resistance”
that is dependant on phytohormone
salicylic acid.
• An attacking pathogen secretes effectors
(Avr) that are recognized by plant
resistance proteins (R) which trigger the
development of hypersensitive response
(HR) which activates a SAR signal.
Protection Response • The SAR signal leads to production of
protection proteins (PR) throughout the
plant which act against a broad spectrum
of the same or other pathogens.
• One of these is the “Shikimic acid
pathway” (phyto alexins) which takes
away nutrients from around the disease
infected areas starving the disease. P is
important.
Shikimic Acid Pathway
Protection Response
• Attacks by predators/diseases trigger
signals that provide defense. In canola
there is a glucosinolate/myrosinase
system triggered that produces more
glucosinolates and trichomes which are
stored in the cell vacuoles to protect the
plant.
Protection Response
• The high levels of glucosinolates may
deter some insects/diseases BUT may also
attract specialist bugs through phenols
that are given off by the plant.
• Sulphur is essential for production of
glucosinolates.
Conclusion
• Understanding precisely how plant
signaling systems and hormones regulate
plant growth is one of the key areas of
fundamental plant biology which will
underpin crop improvements in the
future!
Thank You