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Plant Nutrition – Assimilation and transport

How do plants optimize their uptake and utilization


of nitrogen?

How is inorganic How do plants


nitrogen sense local soil
assimilated into nitrogen levels
organic and plant
molecules? nitrogen status?
How do plants
remobilize
How is nitrogen nitrogen to
taken up into How do plants optimize N-
the plant? respond to nitrogen utilization?
deficit? How do they
maximize uptake
through their roots?
Nitrogen metabolism: Uptake, assimilation and
remobilization
Remobilization Carbon pools
Amino acid Assimilation TCA cycle
Uptake recycling,
Glutamate 2-oxoglutarate
photorespiration
NH4+ Glutamine-2-
oxoglutarate
NH4+ aminotransferase
Nitrite (GOGAT)
reductase
Glutamate
NO2 -
Nitrate
reductase Glutamine Glutamine
synthetase
Incorporation into
NO3- NO3- (GS)
amino acids and
other nitrogen-
R-NH2 containing
NH4+ compounds
Assimilation
N2

Adapted from Xu, G., Fan, X. and Miller, A.J. (2012). Plant nitrogen assimilation and use efficiency. Annu. Rev. Plant Biol. 63: 153-182.
Most plants take up most of their nitrogen as nitrate
NO3-

Many prokaryotes oxidize NH4+, Plants use energy to reduce NO3- for
so soil NH4+ levels are often low assimilation into organic compounds

Energy Energy Energy Energy


released released consumed consumed

NH4+ NO2- NO3- NO3- NO2- NH4+ R-NH3


Nitrate Nitrite
Nitrification by nitrifying prokaryotes reductase reductase

Plant preferences for NH4+ vs NO3- vary by species, other


metabolic processes, temperature, water, soil pH etc….

See Li, B., Li, G., Kronzucker, H.J., Baluška, F. and Shi, W. (2014). Ammonium stress in Arabidopsis: signaling, genetic loci, and physiological targets. Trends
Plant Sci. 19: 107-114; Britto, D.T. and Kronzucker, H.J. (2013). Ecological significance and complexity of N-source preference in plants. Ann. Bot. 112: 957-963.
Plants have specific transporters for NO3-, NH4+ and
other N forms

HATS = high affinity


transporters
LATS = low affinity
transporters

Nacry, P., Bouguyon, E. and Gojon, A. (2013). Nitrogen acquisition by roots: physiological and developmental mechanisms ensuring
plant adaptation to a fluctuating resource. Plant Soil. 370: 1-29, With kind permission from Springer Science and Business Media
A major nitrate importer was the first cloned: CHL1/
NRT1.1/ NPF6.3

The first nitrate Chlorate (ClO3-)


transporter was mimics nitrate Nitrate
identified using a (NO3-) reductase
genetic selection for Chlorite
chlorate resistance ClO2-

Nitrate Chlorate
Wild- In 1993 the CHL1 gene was cloned and
reductase uptake
found to be a nitrate transporter
1973 type mutant mutant
(shown = current in Xenopus oocytes)
(chl1-5)
Growth on
chlorate
Nitrate
reductase
activity
+ - +
Oostindiër-Braaksma, F.J. and Feenstra, W.J. (1973). Isolation and characterization of chlorate-resistant mutants of Arabidopsis thaliana. Mutation Research/Fundamental and Molecular Mechanisms of Mutagenesis. 19: 175-185; Reprinted from Tsay,
Y.-F., Schroeder, J.I., Feldmann, K.A. and Crawford, N.M. (1993). The herbicide sensitivity gene CHL1 of arabidopsis encodes a nitrate-inducible nitrate transporter. Cell. 72: 705-713 with permission from Elsevier.
Other channels contribute to nitrate transport w/in
and between cells

Nitrogen uptake but also assimilation and


recycling depend on membrane transporters

Specific transporters move nitrate


(or other N-containing
compounds) inwards and
outwards across the PM and
across the vacuolar membrane

Reprinted from Wang, Y.-Y., Hsu, P.-K. and Tsay, Y.-F. (2012). Uptake, allocation and signaling of nitrate. Trends Plant Sci. 17: 458-467; Tegeder, M. (2014). Transporters involved in source to sink
partitioning of amino acids and ureides: opportunities for crop improvement. J. Exp. Bot. 65: 1865-1878
Primary N assimilation: NO3- is reduced to NH4+
prior to assimilation

Glutamate
Uptake

NH4+
Assimilation
into organic
Nitrite NH4+ compounds
reductase
NO2-
Nitrate All other N-
reductase Glutamine Glutamine
synthetase containing
NO3- (GS) compounds
NO3-

R-NH3
Nitrate reductase is a large enzyme with a complex
catalytic scheme

NADH NAD+ Nitrate reductase


reduces nitrate to nitrite
NO3- NO2-
with NADH acting as the
electron donor

NO3-
NADH

The electrons move from


NADH to FAD to heme
to a molybdenum
cofactor (Moco) to NO3-

Lambeck, I.C., Fischer-Schrader, K., Niks, D., Roeper, J., Chi, J.-C., Hille, R. and Schwarz, G. (2012). Molecular
mechanism of 14-3-3 protein-mediated inhibition of plant nitrate reductase. J. Biol. Chem. 287: 4562-4571.
GS/GOGAT assimilates inorganic nitrogen into
organic molecules

Remobilization
Carbon pools
Amino acid
Assimilation TCA cycle
recycling,
photorespiration Glutamate 2-oxoglutarate

Glutamine-2-
oxoglutarate
NH4+ aminotransferase
(GOGAT)
Uptake
Glutamate

Glutamine Glutamine
synthetase
Incorporation into
(GS)
amino acids and
other nitrogen-
containing
compounds
Gln synthetase (GS) expression is regulated by many
factors

GS1 (GLN1 genes)


Cytosolic protein

GS2 (GLN2 genes)


Nuclear gene, plastid localized protein

GS activity is regulated transcriptionally


and post-transcriptionally by cell type,
light, [NH4+], circadian cycles, plant
carbon status etc.

GS ac&vity is correlated
with nitrogen use
efficiency

Martin, A., et al., and Hirel, B. (2006). Two cytosolic glutamine synthetase isoforms of maize are specifically involved in the control of grain production. Plant Cell. 18: 3252-3274.
Rebmobilization of N occurs during senescence and
photorespiration

Each N atom may cycle through GS assimilation


many times as amino acids are recycled
during growth and senescence and
remobilization
released due to photorespiration

Leaves Roots, Cotyledons


Amino acids Amino acids remobilization

Glutamate Glutamate
Fdx-GOGAT NADH-GOGAT

Glutamate Glutamine Glutamate Glutamine assimilation


Chloroplast Cytosolic GS1
localized GS2

Photo- NH4+ AA NH4+


respiration catabolism
uptake
Assimilation Assimilation

Avice, J.-C. and Etienne, P. (2014). Leaf senescence and nitrogen remobilization efficiency in oilseed rape (Brassica napus L.). J. Exp. Bot. 65: 3813-3824
In some plants, most grain N is remobilized from
vegetative tissues

The relative amount of N


taken up pre- and post-
flowering is important in
nitrogen use efficiency

Different crop rely more or


less on N remobilization
from vegetative tissues

Hirel, B., Le Gouis, J., Ney, B. and Gallais, A. (2007). The challenge of improving nitrogen use efficiency in crop plants: towards a more central role for
genetic variability and quantitative genetics within integrated approaches. J. Exp. Bot. 58: 2369-2387
Summary: Plant nitrogen uptake and assimilation

Remobilization Carbon pools


Amino acid Assimilation TCA cycle
Uptake recycling,
Glutamate 2-oxoglutarate
photorespiration
NH4+ Glutamine-2-
oxoglutarate
NH4+ aminotransferase
Nitrite (GOGAT)
reductase
Glutamate
NO2 -
Nitrate
reductase Glutamine Glutamine
synthetase
Incorporation into
NO3- NO3- (GS)
amino acids and
other nitrogen-
R-NH2 containing
NH4+ compounds
Assimilation
N2

Adapted from Xu, G., Fan, X. and Miller, A.J. (2012). Plant nitrogen assimilation and use efficiency. Annu. Rev. Plant Biol. 63: 153-182.
Regulation: Nitrogen sensing, signalling and deficit
responses
NITROGEN DEFICIT
Increase uptake
Activation of some NO3- and NH4+ transporters
Preferential growth of root relative to shoot

Metabolic adaptations to low-N


Decreased accumulation of N-rich chlorophyll
Increased accumulation N-free anthocyanins
Smaller pools of N-containing compounds (amino acids)
Larger pools of N-free compounds (starches, organic acids)

Accelerated senescence and nitrogen


remobilization

See for example Scheible, W.-R., et al and Stitt, M. (2004). Genome-wide reprogramming of primary and secondary metabolism, protein synthesis, cellular growth processes, and the regulatory infrastructure of Arabidopsis in response to nitrogen. Plant
Physiol. 136: 2483-2499; Krapp, A. et al and Daniel-Vedele, F. (2011). Arabidopsis roots and shoots show distinct temporal adaptation patterns toward nitrogen starvation. Plant Physiol. 157: 1255-1282. Schlüter, U., et al. and Sonnewald, U. (2012).
Maize source leaf adaptation to nitrogen deficiency affects not only nitrogen and carbon metabolism but also control of phosphate homeostasis. Plant Physiol. 160: 1384-1406. Amiour, N. et al and Hirel, B. (2012). The use of metabolomics integrated
with transcriptomic and proteomic studies for identifying key steps involved in the control of nitrogen metabolism in crops such as maize. J. Exp. Bot. 63: 5017-5033. Balazadeh, S., et al. and Mueller-Roeber, B. (2014). Reversal of senescence by N
resupply to N-starved Arabidopsis thaliana: transcriptomic and metabolomic consequences. J. Exp. Bot. 63: 5017-5033.
Responses to NO3- can be separated from those to
N-metabolites

X
NR mutant can’t grow 10% of the
on NO3- NO3- NO2- NH4+ R-NH3 genome responds
Nitrate Nitrite to nitrate, but only
reductase reductase
some genes are
Nitrate reductase mutants allow responses to NO3- nitrate-specific
to be separated from responses to N-metabolites

Red indicates
nitrate-specific
genes

Transcriptional responses
to nitrate (+ downstream
metabolites)

Wang, R., Tischner, R., Gutiérrez, R.A., Hoffman, M., Xing, X., Chen, M., Coruzzi, G., Crawford, N.M. (2004). Genomic analysis of the nitrate response using a nitrate reductase-null mutant of Arabidopsis. Plant Physiol. 136: 2512–2522; Canales, J.,
Moyano, T.C., Villarroel, E. and Gutiérrez, R.A. (2014). Systems analysis of transcriptome data provides new hypotheses about Arabidopsis root response to nitrate treatments. Front. Plant Sci. 5: 22.
CHL1/NRT1.1/NPF6.3 is a nitrate transceptor
(sensor)

In wild-type Lateral roots of Transceptor mutants


plants (Ws), transceptor mutants (chl1-5) also show
lateral root (chl1-10) fail to abnormal transcripAonal
growth is respond to the HN responses to nitrate
sAmulated in environment
High Nitrate
(HN)

WT

chl1-5

Remans, T., et al. and Gojon, A. (2006). The Arabidopsis NRT1.1 transporter participates in the signaling pathway triggering root colonization of nitrate-rich patches. Proc. Natl. Acad. Sci. 103: 19206-19211 © by the
National Academy of Sciences; Krouk, G., et al. and Gojon, A. (2010). Nitrate-regulated auxin transport by NRT1.1 defines a mechanism for nutrient sensing in plants. Devel. Cell. 18: 927-937
Roots respond to local and systemic nitrogen
availability

When nitrogen is abundant,


When nitrogen
plants allocate less biomass
distribution is patchy,
to their roots
roots proliferate in the
nutrient rich patches

Reprinted by permission from Wiley from Drew, M.C. (1975). Comparison of the effects of a localised supply of phosphate, nitrate and ammonium and potassium on the growth of the seminal
root system, and the shoot, in barley. New Phytol. 75: 479-490.. Reprinted from Bouguyon, E., Gojon, A. and Nacry, P. (2012). Nitrate sensing and signaling in plants. Sem. Cell Devel. Biol. 23:
648-654, with permission from Elsevier. See also Gersani, M. and Sachs, T. (1992). Development correlations between roots in heterogeneous environments. Plant Cell Environ. 15: 463-469.
The split-root system separates local and systemic
signals
All plants split with ½ root system in each of two chambers
Sp.NO3 roots Sp.KCl roots
C.NO3 plants experience locally Experience locally C.KCl plants
Both chambers high NO3- but also deficient NO3- Both chambers
contain KNO3 N-deficiency signals condiAons but also contain KCl (local
(local and systemic derived from Sp.KCl N-sufficient signals and systemic signals
signals indicate roots from Sp.NO3 roots indicate NO3-
NO3 available) deficiency)

Ruffel, S., Krouk, G., Ristova, D., Shasha, D., Birnbaum, K.D. and Coruzzi, G.M. (2011). Nitrogen economics of root foraging: Transitive closure of the nitrate–cytokinin relay and distinct systemic signaling for N
supply vs. demand. Proc. Natl. Acad. Sci. USA 108: 18524-18529.
Evidence for a systemic signal of "
N-demand on root development
Signals from the N- Signals from the N-replete
deficient roots promote Sp.NO3 roots supress root
elevated root growth in growth in Sp.KCl as compared to
Sp.NO3 as compared to C.KCl roots, indicaAng that a
C.NO3, indicating that a response to systemic N-
response to systemic N- reple&on signals
starvation signals

Model: Systemic
signals promote root
growth and suppress
root growth

Ruffel, S., Krouk, G., Ristova, D., Shasha, D., Birnbaum, K.D. and Coruzzi, G.M. (2011). Nitrogen economics of root foraging: Transitive closure of the nitrate–cytokinin relay and distinct systemic signaling for N
supply vs. demand. Proc. Natl. Acad. Sci. USA 108: 18524-18529.
Evidence for cytokinin-dependent and –independent
signals

*
A separate signal that promotes
root growth in plants with total N
deprivation (C.KCl) still operates
in CK-deficient plants, as shown
by increased growth in C.KCl as
compared to Sp.KCl conditions
In cytokinin
deficient plants,
there is no
Growth augmentation
systemic N-
correlating to N-starvation
demand induced
increase in root
length

Ruffel, S., Krouk, G., Ristova, D., Shasha, D., Birnbaum, K.D. and Coruzzi, G.M. (2011). Nitrogen economics of root foraging: Transitive closure of the nitrate–cytokinin relay and distinct systemic signaling for N
supply vs. demand. Proc. Natl. Acad. Sci. USA 108: 18524-18529.
Model of the effects of (some) local and systemic
signals
Systemic Other factors that contribute
Systemic to local and systemic signals
include auxin, amino acids,
transcription factors and root-
derived peptides

Loss-of-funcAon receptor
mutants for root-derived
pepAdes do not
Local NO3 effect downregulate root
growth when N is
abundant

WT LOF

Ruffel, S., Krouk, G., Ristova, D., Shasha, D., Birnbaum, K.D. and Coruzzi, G.M. (2011). Nitrogen economics of root foraging: Transitive closure of the nitrate–cytokinin relay and distinct systemic signaling for N
supply vs. demand. Proc. Natl. Acad. Sci. USA 108: 18524-18529. Guan, P., Wang, R., Nacry, P., Breton, G., Kay, S.A., Pruneda-Paz, J.L., Davani, A., and Crawford, N.M. (2014). Nitrate foraging by Arabidopsis roots is
mediated by the transcription factor TCP20 through the systemic signaling pathway. Proc. Natl. Acad. Sci. USA 111: 15267-15272. Tabata, R., Sumida, K., Yoshii, T., Ohyama, K., Shinohara, H., and Matsubayashi,Y.
(2014). Perception of root-derived peptides by shoot LRR-RKs mediates systemic N-demand signaling. Science 346: 343-346.
Model for transceptor action: NO3- competes for
auxin transport

When NO3- is When NO3- is


low, NPF6.3 high, auxin
transports transport
auxin away through
from the root NPF6.3 is
tip and growth suppressed
is inhibited and growth is
promoted

NPF6.3 Auxin NPF6.3 Auxin


NO3- NO3-

Beeckman, T. and Friml, J. (2010). Nitrate contra auxin: Nutrient sensing by roots. Devel. Cell. 18: 877-878 with permission from Elsevier. See also Krouk, G., et al and Gojon, A. (2010). Nitrate-regulated auxin transport by NRT1.1 defines a
mechanism for nutrient sensing in plants. Devel. Cell. 18: 927-937; Mounier, E., et al and Nacry, P. (2014). Auxin-mediated nitrate signalling by NRT1.1 participates in the adaptive response of Arabidopsis root architecture to the spatial heterogeneity
of nitrate availability. Plant Cell Environ. 37: 162-174; Forde, B.G. (2014). Nitrogen signalling pathways shaping root system architecture: an update. Curr. Opin. Plant Biol. 21: 30-36.
Strategies to improve nitrogen-use efficiency and
decrease N pollution
Nitrogen fixation is
energy demanding
Nitrous oxide (N2O)
N N O derived from fertilizer is a
major greenhouse gas

Cyanabacterial bloom

Lake Erie Unhealthful nitrate from agricultural


uses pollutes groundwater

Nolan, B.T. and Hitt, K.J. (2006). Vulnerability of shallow groundwater and drinking-water wells to nitrate in the United
States. Environ. Sci. Technol. 40: 7834-7840. Image source: Lamiot; Alexandra Pugachevsky; NASA Earth Observatory
Co-cropping and monitoring can decrease the need
for N application

Co-cropping or
growing in rotation
with legumes enriches
soil N content

N status can be Chlorophyll can


determined by be measured
chlorophyll content, by the
measured by transmission
reflected light ratio of 653 nm
to 931 nm light

Apogee; N2Africa; Petr Kosina / CIMMYT. See also Muñoz-Huerta, R.F., Guevara-Gonzalez, R.G., Contreras-Medina, L.M., Torres-Pacheco, I., Prado-Olivarez, J., and
Ocampo-Velazquez, R.V. (2013). A review of methods for sensing the nitrogen status in plants: Advantages, disadvantages and recent advances. Sensors. 13:
10823-10843; Robertson, G.P. and Vitousek, P.M. (2009). Nitrogen in agriculture: Balancing the cost of an essential resource. Annu. Rev. of Environ. Res. 34: 97-125.
Slow-release fertilizers can match release to
requirements

Traditional fertilizers don’t match


Traditional fertilizer –
nitrogen availability to plant needs.
one or two applications Slow release fertilizers can more
closely match plant needs
Amount of fer9lizer available

Slow-release
fer9lizer
Time
H 2O N N
UREA
H 2O N

Plant growth
requirements Coated urea dissolves and releases
slowly, but it can be expensive
Time

Adapted from Timilsena, Y.P., Adhikari, R., Casey, P., Muster, T., Gill, H. and Adhikari, B. (2014). Enhanced efficiency fertilisers: a review of formulation and nutrient release patterns. J. Sci. Food Agric. DOI: 10.1002/jsfa.6812
Soil bacteria can be manipulated to decrease N2O
and NO3- pollution

Inhibitors of bacterial nitrification cause


NH4+ to be retained in the soil, leading to
less leaching and less N2O production

Denitrifying bacteria cultivated in


a bioreactor downstream of a
fertilized field protect waterways
by converting NO3- in runoff to N2

Philippot, L. and Hallin, S. (2011). Towards food, feed and energy crops mitigating climate change. Trends Plant Sci. 16: 476-480 with permission from Elsevier. See also Subbarao, G.V., et al. 2009). Evidence for biological nitrification inhibition in
Brachiaria pastures. Proc. Natl. Acad. Sci. USA. 106: 17302-17307. Subbarao, G.V., et al., (2013). A paradigm shift towards low-nitrifying production systems: the role of biological nitrification inhibition (BNI). Ann. Bot. 112: 297-316; Schipper,
L.A., Robertson, W.D., Gold, A.J., Jaynes, D.B. and Cameron, S.C. (2010). Denitrifying bioreactors—An approach for reducing nitrate loads to receiving waters. Ecol. Engin. 36: 1532-1543.
Altering flux into amino acid pools can increase NUE

Remobilization
Carbon pools
Amino acid Storage
Assimilation TCA cycle
recycling,
photorespiration Glutamate 2-oxoglutarate
Alanine
Glutamine-2-
Alanine
oxoglutarate
aminotransferase
NH4+ aminotransferase
(AlaAT)
(GOGAT)
Uptake
Glutamate Pyruvate

Glutamine Glutamine
synthetase
Incorporation into
(GS)
amino acids and
other nitrogen-
containing
compounds

Good, A.G., Johnson, S.J., De Pauw, M., Carroll, R.T., Savidov, N., Vidmar, J., Lu, Z., Taylor, G. and Stroeher, V. (2007). Engineering nitrogen use efficiency with alanine aminotransferase. Can. J. Bot. 85: 252-262.
Breeding strategies for enhanced nitrogen use
efficiency
Glutamine synthetase
Traits of an acJvity is an important
idealized plant component of NUE
with high NUE

In rice, a subunit of a
heterotrimeric G
protein contributes
to N-sensiJve growth
and N assimilaJon

Chardon, F., Noël, V. and Masclaux-Daubresse, C. (2012). Exploring NUE in crops and in Arabidopsis ideotypes to improve yield and seed quality. J. Exp. Bot. 63: 3401-3412 by permission of Oxford University Press;
Martin, A., et al. and Hirel, B. (2006). Two cytosolic glutamine synthetase isoforms of maize are specifically involved in the control of grain production. Plant Cell. 18: 3252-3274. Reprinted by permission from
Macmillan Publishers Ltd: Sun, H., et al. (2014). Heterotrimeric G proteins regulate nitrogen-use efficiency in rice. Nat Genet. 46: 652-656. Hirel, B., Le Gouis, J., Ney, B. and Gallais, A. (2007). The challenge of
improving nitrogen use efficiency in crop plants: towards a more central role for genetic variability and quantitative genetics within integrated approaches. J. Exp. Bot. 58: 2369-2387
Summary: Improving N use efficiency in plants and
soils

•  N is abundant as N2, but often limiting for growth


•  N is fixed by biological or industrial means
•  N fertilization is economically and environmentally
costly
•  N use efficiency involves uptake of NO3- and NH4+,
primary assimilation and recycling via GS / GOGAT
•  Regulatory and signaling pathways are being identified
as opportunities for breeding improvements
•  Monitoring of plant and soil N status can improve
fertilizer use efficiency
Phosphorus (P) is assimilated and used
as phosphate (Pi) which depending on
the pH is H2PO4- ,HPO42- or PO43-
PHT1 phosphate transporters mediate uptake and
transport

9 PHT1 genes in
Arabidopsis, 13 in rice,
Most are 12 in poplar. Some are
expressed in mycorrhiza inducible
roots and other
tissues

PHT transporters are H+/


PO43- co-transporters
that have 12 membrane-
spanning domains

Nussaume, L., Kanno, S., Javot, H., Marin, E., Pochon, N., Ayadi, A., Nakanishi, T.M. and Thibaud, M.-C. (2011) Phosphate import in plants: focus on the
PHT1 transporters. Front. Plant Sci. 2: 83. Pedersen, B.P., et al and and Stroud, R.M. (2013). Crystal structure of a eukaryotic phosphate transporter. Nature.
496: 533-536. Loth-Pereda, V.,et al. and Martin, F. (2011). Structure and expression profile of the phosphate Pht1 transporter gene family in mycorrhizal
Populus trichocarpa. Plant Physiol. 156: 2141-2154. See also Lapis-Gaza, H.R., Jost, R., and Patrick M Finnegan, P.M. (2014). Arabidopsis PHOSPHATE
TRANSPORTER1 genes PHT1;8 and PHT1;9 are involved in root-to-shoot translocation of orthophosphate. BMC Plant Biol. 14: 334.
P-Starvation Inducible responses increase P uptake
and recycling

Proteaceae show metabolic adaptions to P-


impoverished soils such as very efficient use of P

Ribosomes (rRNA) are the major form of organic P.


Proteaceae maintain a very low copy number of
ribosomes, yet are photosynthetically efficient

Proteaceae also show delayed greening;


ribosomes first promote growth, then
chloroplast matura9on

Huang, T.-K., et al and Lucas, W.J. (2014). Molecular mechanisms underlying phosphate sensing, signaling, and adaptation in plants. J. Integr. Plant Biol. 56: 192-220 by permission. Sulpice, R., et al and Lambers, H.
(2014). Low levels of ribosomal RNA partly account for the very high photosynthetic phosphorus-use efficiency of Proteaceae species. Plant Cell Environ. 37: 1276-1298. See also Lin, W.-Y., Huang, T.-K., Leong, S.J.
and Chiou, T.-J. (2014). Long-distance call from phosphate: systemic regulation of phosphate starvation responses. J. Exp. Bot. 65: 1817-1827.
PSI (phosphate-starvation induced) are up-regulated
by PHR1

PSI genes encode


phosphatases,
transporters,
regulatory factors….

The interaction between SPX1


SPX1 interferes with PHR1 and PHR1 is Pi-dependent….
binding to its DNA binding site
(P1BS). In yeast, SPX1 proteins
act as Pi sensors

Puga, M.I., Mateos, I., Charukesi, R., Wang, Z., Franco-Zorrilla, J.M., de Lorenzo, L., Irigoyen, M.L., Masiero, S., Bustos, R., Rodríguez, J., Leyva, A., Rubio, V., Sommer, H. and Paz-Ares, J. (2014). SPX1 is a phosphate-dependent inhibitor of
PHOSPHATE STARVATION RESPONSE 1 in Arabidopsis. Proc. Natl. Acad. Sci. USA 111: 14947-14952; Wang, Z., Ruan, W., Shi, J., Zhang, L., Xiang, D., Yang, C., Li, C., Wu, Z., Liu, Y., Yu, Y., Shou, H., Mo, X., Mao, C. and Wu, P. (2014). Rice
SPX1 and SPX2 inhibit phosphate starvation responses through interacting with PHR2 in a phosphate-dependent manner. Proc. Natl. Acad. Sci. USA 111: 14953-14958.
Regulatory controls prevent Pi from over
accumulating
PHO1 is a transporter that PHO2 is an E2 ligase
moves Pi into xylem for In pho1 mutants, too
that targets much Pi accumulates
transport to the shoot
PHT transporters take up Pi
transporters for in the root and too
proteolysis little in the shoot
Pi In pho2 mutants,
shoot
too much Pi
accumulates in the
shoot and too little
in the root; transport
xylem

is out-of-control

PHO2
root
PHO1 PHO1
PHT Too much
or too little
Pi is bad

Delhaize, E., and Randall, P.J. (1995). Characterization of a phosphate-accumulator mutant of Arabidopsis thaliana. Plant Physiol. 107: 207 – 213; Liu, T.-Y., Huang, T.-K., Tsenga, C.-Y., Lai, Y.-S., Lin, S.-I., Lin,
W.-Y., Chen, J.-W., Chiou, T.J. (2012). PHO2-dependent degradation of PHO1 modulates phosphate homeostasis in Arabidopsis. Plant Cell 24: 2167 – 2183.
Mutants pho1 and pho2 show effects of altered Pi
transport

Liu, T.-Y., Lin, W.-Y., Huang, T.-K. and Chiou, T.-J. MicroRNA-mediated surveillance of phosphate transporters on the move. Trends Plant Sci. 19: 647-655
PHO2 accumulation is regulated by miR399
expression

When Pi is ample, PHO2 P starvation induces miR399


targets PHO1 for expression of miR399,
degradation which targets PHO2 mRNA PHO2
for degradation
A target mimic IPS1
+ Pi - Pi fine-tunes the effects
Pi of miR299; by binding
stably to miR399,
PHO2 PHO2
mRNA mRNA IPS1 supports PHO2
expression
PHO2

xylem
PHO2 PHO2
PHO1 PHO1 miR399
Pi
IPS1

Redrawn from Franco-Zorrilla, J. M., Valli, A., Todesco, M., Mateos, I., Puga, M.I., Rubio-Somoza, I., Leyva, A., Weigel, D., García, J.A.,
and Paz-Ares, J. (2007) Target mimicry provides a new mechanism for regulation of microRNA activity. Nat. Genet. 39: 1033–1037.
P uptake & transport are regulated by local and
systemic signals
Suppression of shoot
branching

Strigolactones
Establishment of
plant – mycorrhizal
fungi symbiosis

Phosphate Enhanced
starvation PHT1 transporters uptake
signal
(unknown) PHR1 Phosphatases,
(transcription factor) organic acid
synthases
PHO1 (miR399 is a negaAve
regulator of a negaAve
IPS1 miR399 PHO2 PHT1 regulator of P uptake)

Wu, P., Shou, H., Xu, G. and Lian, X. (2013). Improvement of phosphorus efficiency in rice on the basis of understanding phosphate signaling and homeostasis. Curr. Opin. Plant Biol. 16: 205-212. Liu, T.-Y., Lin, W.-Y.,
Huang, T.-K. and Chiou, T.-J. (2014). MicroRNA-mediated surveillance of phosphate transporters on the move. Trends Plant Sci. 19: 647-655.
The auxin transport capacity in the primary shoot governs the
induction of shoot branching
An intimate crosstalk of different plant hormones regulates
shoot branching
Strategies to improve crop plant phosphorus use
efficiency

Vinod, K.K. and Heuer, S. (2012). Approaches towards nitrogen- and phosphorus-efficient rice. AoB Plants. 2012: pls028
Selection for root architecture traits can lead to
increased P uptake
P-uptake efficiency can be correlated P-efficient
to more efficient root traits root system

P-inefficient
root system

P-efficient root system P-inefficient


root system

Lynch, J.P. (2011). Root phenes for enhanced soil exploration and phosphorus acquisition: Tools for future crops. Plant Physiol. 156: 1041-1049; Wang, X., Yan, X. and
Liao, H. (2010). Genetic improvement for phosphorus efficiency in soybean: a radical approach. Ann. Bot. 106: 215-222.
Rice adapted to poor-soil regions revealed a key
protein kinase

•  The Pup1 (Phosphate Uptake 1)


major QTL was identified in aus-
variety rice adapted to poor soils
•  Eventually this was revealed to
encode a protein kinase PSTOL1
not present in other rice genomes
•  Overexpression of PSTOL1 leads
to enhanced root growth
Overexpressor Control

Reprinted by permission from Macmillan Publishers Ltd : Gamuyao, R., Chin, J.H., Pariasca-Tanaka, J., Pesaresi, P., Catausan, S., Dalid, C., Slamet-Loedin, I., Tecson-Mendoza, E.M., Wissuwa, M. and Heuer, S. (2012).
The protein kinase Pstol1 from traditional rice confers tolerance of phosphorus deficiency. Nature. 488: 535-539.See also Chin, J.H., Gamuyao, R., Dalid, C., Bustamam, M., Prasetiyono, J., Moeljopawiro, S., Wissuwa,
M. and Heuer, S. (2011). Developing rice with high yield under phosphorus deficiency: Pup1 sequence to application. Plant Physiol. 156: 1202-1216.
Is it feasible to reuse, recapture and recycle
phosphate?
Phosphate recovered
from human urine alone Human urine is rich
in phosphate, and it Cleaner

P and N can be precipitated out of wastewater


could replace >20% of
wastewater
phosphate demands can be separated out
from other waste at
the point of origin

Mg

Mg
Urine can be
applied directly to
plants as liquid
fertilizer

N & P-rich
Wastewater in Struvite
P (NH MgPO ·6H O)
P crystals harvested
for use as fertilizer
Urine-reclaiming toilet

Pratt, C., Parsons, S.A., Soares, A. and Martin, B.D. (2012). Biologically and chemically mediated adsorption and precipitation of phosphorus from wastewater. Curr. Opin. Biotechnol. 23: 890-896; Mihelcic,
J.R., Fry, L.M. and Shaw, R. (2011). Global potential of phosphorus recovery from human urine and feces. Chemosphere. 84: 832-839.. Multiformharvest.com
Strategies have been developed to impede P from
entering waterways

Chemical and biological processes


including algal production can
effectively remove P from wastewaters

Timing of fertilizer application and


management of water flow from can decrease
the amount of P that enters waterways

McDowell, R.W. (2012). Minimising phosphorus losses from the soil matrix. Curr. Opin. Biotech. 23: 860-865 with permission from Elsevier; Pratt, C., Parsons, S.A., Soares, A. and Martin, B.D. (2012). Biologically
and chemically mediated adsorption and precipitation of phosphorus from wastewater. Curr. Opin. Biotech. 23: 890-896 Shilton, A.N., Powell, N. and Guieysse, B. (2012). Plant based phosphorus recovery from
wastewater via algae and macrophytes. Curr. Opin. Biotech. 23: 884-889 by permission from Elsevier, and others from the same issue. Rittmann, B.E., Mayer, B., Westerhoff, P. and Edwards, M. (2011). Capturing the
lost phosphorus. Chemosphere. 84: 846-853. Schipper, W. (2014). Phosphorus: Too big to fail. Eur. J. Inorgan. Chem. 2014: 1567-1571.
Summary: Phosphorus

•  First or second most commonly limiting nutrient


•  Very insoluble and immobile in soil
•  Roots mine and forage for P through exudations and
symbioses
•  Root system architecture is particularly sensitive to P
•  Uptake involves positive and negative controls
•  Strategies are available to minimize P pollution

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