Life on earth depends on water
Transcript of Life on earth depends on water
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Life on earth depends on water
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Osmosis –
diffusion through semipermeable membrane
Diffusion
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Aquaporins
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Water uptakeCapilary Kapilarwasser – Wurzelhaare – Wurzel – Leitgewebe – Blätter
Quellung der Wurzelwand
apoplastischer Transport versus symplastischer Transport
Aufnahme ins Cytoplasma der Wurzelhaare
Verteilung in Rhizodermisgewebe
Casparischer Streifen
Endodermis – Zentralzylinder
z. T. aktiver Transport, Aquaporine, Oocytenexperiment
Wurzeldruck (Dekapitierungsexperiment)
Xylemtransport, Kohäsion, Adhäsion
Max. Baumhöhe 130 m
Verletzung des Xylems: Reparatur oder Toxizität
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Suberin-Einlagerung im Casparischen Streifen
(in der Endodermis)
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Water transport is based on …
Root Pressure osmotic pressure within
the root cells that causes sap to rise
Transpirationcuticular (10%) stomata (90%)
Increase in osmotic pressure: stomata opening
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Potometer
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Ions
growth, storage (e.g. vacuole), complex formation, secretion (cell wall)
macro-, microelements
active/passive uptake
essentail ions: e.g.
Ca, Na, Fe, Mn, Zn, Cu, NO3-, Mg, PO4
3-, SO42-, B, Mb
toxic heavy metal ions
Cd, Hg, etc.
control of uptake:
concentration gradients, transporter
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Transport mechanisms
01
2
3
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Electrochemical equilibrium develops in which the Na+ concentration gradient, indicated by the upper arrow, is just balanced by the elctrical
gradient, indicated by the lower arrow. At this point, there is no net flux of Na+ to either side.
Electrochemical gradients
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Patch clamp
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Identification and characterization of plant transporters by yeast complementation
Yeast transporter mutantTransformation with plant cDNA library
Test after transformation into plant
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Heavy metal uptake and detoxification
Cd stress
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40 metals
6 g/cm3
- Cd, Cr, Cu, Pb, Ni, Tl, Hg, Zn, As, B, Fe, Mo, Mn, Zn-Fe, Mo, Mn, Zn = essentiel trace elements
- phytoremediation(e.g. with transgenic plants)
Resistence strategies:a. No uptake
b. Storage (vacuole) c. secretion (cell wall)d. Complex formation
(metallothionein, phytochelatin, organic compounds)
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Resistant plants
a. Excluder
b. Indikator
c. Accumulator
Transporters are not always specific
(co-transport of Cd, Zn, Co)
Cu plant from Sambia: Cu-accumulator (+ resistence against As)
Arabidopsis halleri: Cd-accumulator
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- plants and animals
- small proteins with many cyctein residues
- cytoplasma
- complex free metal ions
Metallothionin
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Peptide with 1-6 repetitive gamma-glutamyl-cysteinyl units
- SH groups of cystein bind heavy metals as thiolates
- preference for Cd, Cu, Pb, (Zn)
-
Phytochelatin (PC) synthase
Transfers Glu-Cys-rest onto the tripeptid glutathione
Interesting: posttranslational regulation of PC synthese enzyme
Heavy metals activate PC-synthase activity
Arabidopsis PC-synthase ovrexpressor: heavy metal resistance
Phytochelatin
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Roots are associated with beneficialmicrobes
Mycorrhizal fungi
beneficial endphytic fungi and bacteria
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Beneficial interaction between plants and microbes
N2fixing bacteria/legumes
nodule “
mycorrhiza
endomycorrhizaectomycorrhiza
“
endophytes
bacteriafungi
roottip “
“
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Common feature:Promotion of plant performance
1. Colonisation of roots 2. Promotion of growth & biomass
3. Resistanz aginst abiotic & biotic stress
drought heavy metal pathogen infection
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mycorrhial fungi and rhizobacteriafew host plants
endophytesoften colonizes many hosts
important for agricultureeasier recognition mechanisms
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Pi-504 - CellotrioseC H O
biomolecules
Myc-factor (LCO#)C H O N S
Nod-factor (LCO#)C H O N S
P. indica (Sebacinales)
host range
AM fungi (Glomeromycota)
Rhizobia (Rhizobiales)
all planta tested so far
P. indica
AM fungus*
Rhizobium*
microbe
moss* monocots A. thaliana* dicots
monocots dicots
legumes
[Ca]
2+cy
t(µ
M)
[Ca]
2+cy
t(µ
M)
[Ca]
2+cy
t(µ
M)
#LCO: lipochitooligosaccharide
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Both partners MUST profit from the symbiosis.
Photosynthesis“
sucrosewater, soil nutrients,
phosphate, nitrogen
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Two types of mycorrhiza
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Beneficial interaction between plants and microbes
N2 fixingbacteria/legumes
nodule “
mycorrhiza
endomycorrhizaectomycorrhiza
“
endophytes
bacteriafungi
roottip “
“
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Endomycorrhiza
arbuscles
Fungal and plant cells areCOMPLETELY separated.
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- 80% of endomycorrhizal fungi are arbuscular mycorrhiza (VAM)
- only 6 fungal species [Glomales, Zygomycetes] form VAM
Initiation:
- germination of spores- Hyphae form appressorium
(attachment sites)
- Formation of an extracellular hyphal system in the apoplast
Endomycorrhiza
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Penetration:
- formation of haustorium: penetration into the plant cell
- Extracellular hyphae:
- collection of nutrients from rhizosphere
- Intracellular hyphae:
- enlargement of interaction surface
- nutrient/metabolite exchange with plant cell
Life time of arbuscle: a few days
extracellularhyphae
intracellular arbuscles
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Ektomycorrhiza
hyphae: rhizosphere, on root surface, between root cellsdo not enter vascular tissue
fruit bodies
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• many trees form ectomycorrhizas
• Hartig Net:– a net around the root (hairs) to extent access to soil nutrients
Hartig Net
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• Optimization of nutrient exchanges
• Protection against soil pathogens
• Connection of organisms in biophere– 60 km network– Support for young trees in forests– Connections between different plant species
Function of Hartig Net
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Beneficial interaction between plants and microbes
N2 fixingbacteria/legumes
nodule “
mycorrhiza
endomycorrhizaectomycorrhiza
“
endophytes
bacteriafungi
roottip “
“
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Nodules
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• Haber-Bosch: N-fertilizer
• N2 fixation – rhizobacteria– cyanobacteria
N2 + 8H+ + 8e- + 16 ATP = 2NH3 + H2 + 16ADP + 16 Pi
N2 fixation
rhizobia withnitrogenase
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bacterial nitrogenase
Ferox
Ferred
ATP
ADP
NH4+
N2
photosynthesis
- nitrogenase O2-sensitive
- leghaemoglobin (plant- and bacteria-encoded)
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Rhizobia under N limitation
alfalfa soybean
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recognition
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Endophytes
GOALS:
- Isolation of endophytic fungi from roots of plants growing under extreme conditions(deserts, salt lakes, high mountains, flooded areas)
- Cultivation of fungi, identification- Test with model plant Arabidopsis thaliana under extreme conditions- Molecular mechanisms of resistance
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Piriformospora indica - plant growth promoting fungus
+ Pi- Pi
N. tabaccum
4 weeks
+ Pi
- Pi
10 days
Pi colonized Arabidopsis root
Kaefer medium
KM broth
Oelmüller et al. (2009) Symbiosis, 49, 1-17; Varma et al. (2012) Agric Res, 1, 117-131. Johnson et al. (2011) Endocyt Cell Res, 21, 101-113.
7 weeks
- Pi + Pi
- Pi
4 weeks
+ Pi
- PiA. thaliana Pi: P. indica
+ Pi
- Pi
10 days
Pi: P. indica2
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Piriformospora indica – lateral root development
- fungus + fungus
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Luminometer
Recognition by [Ca2+]cyt –inducing chemical mediators
Aequorea victoria
[Ca2+]apo
[Ca2+]cyt
Channels
Apoaequorin transformed plants
+
Coelenterazine & O2
Aequorin [Ca2+]cyt
Blue light (»=469 nm)
7
gene from
Chemical mediator from
fungusRecognition by host and
signaling opens Ca2+ channels
Transgenic root cell
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e.g. cellotriose
[Ca]
2+cy
t(µ
M)
0,0
0,1
0,2
0,3
0,4
0,5
0,6
0 2 4 6 8 10
Chemcial mediator
water
Time (min)
9