Plants are ~70 to

>90% water by
weight N
Nitrogen

P Phosphorus
K
CO2, Ca Calcium
photo- Potassium
Mg Magnesium
synthesis
S Sulfur
42% Carbon 7% Other,
Si Silicon
from soil Cl Chlorine
Other
7% Hydrogen
These elements are
44% Oxygen obtained mainly from soil,
93% of plant are often referred to as
dry mass is
mineral nutrients, and are
composed of H2O water
C, O and H the subject of the topic
Plant Nutrition
 Plants assimilate mineral nutrients
from their surroundings

Nutrient assimilation is very

energetically demanding –
the nutrients have to be
moved against a
concentration gradient and
often a charge gradient

Tracheophytes
K+
Algae NO3-
NO3-
NO3-

K+ K+

PO43- Bryophytes K+
K+
K+ PO43-
PO4 3-
PO43- PO43-
-
PO43-
NO3

K+ K+
 Plants assimilate mineral nutrients
mainly as cations or anions
MACRONUTRIENTS MICRONUTRIENTS
μmol / g Element Assimilated μmol / g Element Assimilated
(dry wt) form (dry wt) form
250 Potassium (K) K+ 2 Iron (Fe) Fe3+, Fe2+
1000 Nitrogen (N) NO3-, NH4+ 0.002 Nickel (Ni) Ni2+
60 Phosphorus HPO42-, 1 Manganese Mn2+
(P) H2PO4- (Mn)
30 Sulfur (S) SO42- 0.1 Copper (Cu) Cu2+
80 Magnesium Mg2+ 0.001 Molybdenum MoO42+
(Mg) (Mo)
125 Calcium (Ca) Ca2+ 2 Boron (B) H3BO3
3 Chlorine (Cl) Cl-
0.3 Zinc (Zn) Zn2+
See Taiz, L. and Zeiger, E. (2010) Plant Physiology. Sinauer Associates; Marschner, P. (2012) Mineral Nutrition of Higher Plants. Academic Press, London
K + dan Na + - “Si kembar”. Begitu
mirip namun sangat berbeda
Potassium NaCl
deficiency toxicity
Natrium (Na) dan Kalium (K):
• Kolom yang sama dari tabel
periodik
• Keduanya memiliki satu elektron di
kulit terluarnya sehingga
membentuk kation monovalen
• Keduanya merupakan elemen yang
sangat berlimpah

K
Namun, kalium adalah
nutrisi penting, dan natrium
sering kali beracun

Benito, B., Haro, R., Amtmann, A., Cuin, T.A. and Dreyer, I. (2014).The twins K+ and Na+ in plants. J. Plant Physiol. 171: 723–731. FAO
 Penyerapan kalium, transportasi dan
homeostasis
Meningkatkan
kesuburan Mempertahankan
turgor dan
Merangsang mengurangi layu
toleransi stres Mengatur
konduktansi
Mengatur Symptoms of
stomata,
aktivitas enzim potassium deficiency
fotosintesis, dan
transpirasi
Memperkuat
dinding sel
Mempertahankan
Merangsang homeostasis ionik
translokasi
[K+] in soil = ~0.1 – 1 mM
fotosintat [K+] in plant cell
cytoplasm = ~100 mM

See Wang, M., Zheng, Q., Shen, Q. and Guo, S. (2013). The critical role of potassium in plant stress response. Intl. J. Mol. Sci. 14: 7370-7390; Sin Chee Tham /Photo; Purdue extension; Onsemeliot.
 Pupuk kalium ditambang dari bawah
tanah dikenal dengan "kalium/potash”
Potash adalah istilah yang mencakup
banyak bentuk kalium : Hampir setengah dari sumber kalium dunia
• KCl (kalium klorida/silvit) ditemukan di Saskatchewan, Kanada
• K2SO4 (kalium sulfat)
• K2CO3 (kalium karbonat)
• K2Ca2Mg(SO4)4·2H2O (polyhalite)
• etc.

KCl, silvit
Canada Potash; Lmbuga
 Potash menyediakan K+ untuk pupuk,
yang melengkapi sumber alami
Air dipompa
Air dengan
ke bawah
garam K+
tanah
terlarut Aplikasi
kembali ke pupuk kalium
permukaan

0,1 - 0,2%
larutan tanah manure
Garam
decomposition
ditemukan K+
melalui
penguapan 1 - 3% garam
yang dapat Terrestrial
ditukar cycle: Plant /
Animal / Soil
Sumber bawah tanah
90 - 98%
mineral
tidak larut
Adapted from International Potash Institute
 Kalium merupakan unsur hara
esensial
K+ adalah ion pengganti untuk
molekul bermuatan negatif
termasuk DNA dan protein

K+ bergerak masuk
dan keluar dari
vakuola melalui
transporter tertentu

Sebagai kation utama

dalam vakuola, K+
memberikan
kontribusi untuk
ekspansi sel dan
K+ uptake K+ adalah kofaktor gerakan, termasuk
involves high untuk beberapa yang dari sel penjaga
and low affinity enzim
transporters
Reprinted from Maathuis, F.J.M. (2009). Physiological functions of mineral macronutrients. Curr. Opin. Plant Biol. 12: 250-258 with permission from Elsevier.
 Studi awal serapan kalium pada
tanaman: penyerapan Biphasic
Baru-baru ini mutan Arabidopsis
menunjukkan HAK5 dan AKT1
menengahi serapan afinitas tinggi dan
rendah, masing-masing. Data ini
merupakan kesimpulan dari studi mutan.

High affinity Low affinity

transport transport

KCl (mM)

Epstein et al
menunjukkan dua fase
serapan K+ di akar
barley
Epstein, E., Rains, D.W., and Elzam, O.E. (1963). Resolution of dual mechanisms of potassium absorption by barley roots. Proc. Natl. Acad. Sci. USA. 49: 684 – 692;
Gierth, M. and Mäser, P. (2007). Potassium transporters in plants – Involvement in K+ acquisition, redistribution and homeostasis. FEBS Lett. 581: 2348-2356.
Lebih banyak energi harus dikeluarkan untuk
mengambil K + ketika kondisi langka
K+ H+ 2 x ATP ATP
K+
Pengambilan K+ uptake
dari [K+]ext rendah
membutuhkan lebih
banyak energi daripada
2 x H+ H+ ketika [K+]ext lebih tinggi
High affinity Low affinity
transport transport

Pada saat [K+]extsangat tinggi,

non-selective cation channels
(NSCC) juga dapat
berkontribusi

See Britto, D.T. and Kronzucker, H.J. (2008). Cellular mechanisms of potassium transport in plants. Physiol. Plant. 133: 637-650; Nieves-Cordones, M., Alemán, F.,
Martínez, V. and Rubio, F. (2014). K+ uptake in plant roots. The systems involved, their regulation and parallels in other organisms. J. Plant Physiol. 171: 688–695.
 Sel penjaga adalah sistem model
untuk studi transportasi K+
OPEN CLOSING

V-ATPase

H+
H+ aktivitas metabolisme sel
V-PPase penjaga melibatkan ion
A- bergerak masuk dan
H+ H+
A- keluar melintasi
membran plasma dan
TPK
vakuola untuk
H+
mendorong aliran
K+ osmotik air dan
K+
H+ K+ karenanya mengubah
TPC turgor dan ukuran sel
PM-H+-ATPase GORK
KAT K+
Hills, A., Chen, Z.-H., Amtmann, A., Blatt, M.R. and Lew, V.L. (2012). OnGuard, a computational platform for quantitative kinetic modeling of guard cell physiology. Plant Physiol. 159: 1026-1042 Chen, Z.-H., Hills, A.,
Bätz, U., Amtmann, A., Lew, V.L. and Blatt, M.R. (2012). Systems dynamic modeling of the stomatal guard cell predicts emergent behaviors in transport, signaling, and volume control. Plant Physiology. 159: 1235-1251.
Homeostasis kalium: Respons terhadap
ketersediaan K+ yang rendah
Low K

Membrane Hormonal changes

hyperpolarization (auxin, ethylene)

Calcium Enhanced root

Direct effects
signaling growth and
Indirect effects
gravitropic
Transcriptional responses
More efficient
induction of HAK5
uptake through
K+ channel
K+ channels

K+ uptake
Adapted from Chérel, I., Lefoulon, C., Boeglin, M. and Sentenac, H. (2013). Molecular mechanisms
involved in plant adaptation to low K+ availability. J. Exp. Bot. 65: 833-848.
 Toksisitas natrium, transportasi dan
toleransi

You can’t take salt

out of soil easily;
once it is there it
To demonstrate his (fake) stays there
madness, Odysseus plowed
salt into his field
Colum, P. (1918). The Adventures of Odysseus and the Tale of Troy. Project Gutenberg; USDA, USDA, Peggy Greb; FAO
Di daratan, banyak tanah berada di atas endapan
garam kuno yang bisa bergerak ke atas
Pembukaan vegetasi asli sering menyebabkan salinisasi tanah

Salty Salty
water water
Salty water

Department of Agriculture and Rural Affairs (1980). ‘Managing Salinity: Ensuring a Farming Future’. The State of Victoria
 Irigasi juga berkontribusi terhadap salinitas
tanah dengan memobilisasi garam

Rain Rain
Tanpa irigasi, air hujan
Evaporation
tidak menembus di
bawah zona akar

Irigasi berlebihan
menembus ke tanah yang
lebih dalam dan asin,
melarutkan garam dan
menariknya ke atas ke
dalam zona akar
Salt
(dissolved)
 Bagaimana kita bisa mengatasi masalah yang
disebabkan oleh salinisasi tanah?
Hindari menambah
masalah dengan Pelajari tentang
pengelolaan sistem toleransi garam
tanah rapuh yang lebih dari spesies
baik toleran garam
Areas of concern alami (halofit)
Salicornia europaea
Identifikasi halofit yang
Arthrocnemum
dapat digunakan macrostachyum
sebagai tanaman
pangan atau energi

Identifikasi respons
Chenopodium Thinopyrum terhadap stres
quinoa ponticum garam pada
Pelajari kerabat toleran spesies yang
sensitif terhadap
garam pada tanaman
garam (glikofit)
Memperkenalkan sifat toleransi
Munns, R., James, R.A., Xu, B., Athman, A., Conn, S.J., Jordans, C.,
Byrt, C.S., Hare, R.A., Tyerman, S.D., Tester, M., Plett, D. and
salinitas pada tanaman pertanian Geng, Y., Wu, R., Wee, C.W., Xie, F., Wei, X., Chan, P.M.Y.,
Gilliham, M. (2012). Wheat grain yield on saline soils is improved by
an ancestral Na+ transporter gene. Nat Biotech. 30: 360-364.
melalui pemuliaan dan rekayasa Tham, C., Duan, L. and Dinneny, J.R. (2013). A spatio-temporal
understanding of growth regulation during the salt stress response
CSIRO; The State of Victoria; Maurice Chédel; Marco Schmidt in Arabidopsis. Plant Cell. 25: 2132-2154.
 Tumbuhan memiliki toleransi
salinitas yang luas

Saltbush (Atriplex amnicola)

adalah halofit yang dapat
mentolerir tanah yang sangat
asin

Arabidopsis
dan padi
cukup sensitif
Reprinted by permission of Annual Reviews from Munns, R. and Tester, M. (2008). Mechanisms of salinity tolerance. Annu. Rev. Plant Biol. 59: 651-681.
 Mechanisms of sodium toxicity and
tolerance
SALINITY STRESS

Ionic stress: Osmotic stress

K+ deficiency /
excess Na+ influx Oxidative
stress Inhibition of:
water uptake,
Inhibition of:
growth,
enzyme activity,
Detoxification photosynthesis
protein synthesis,
photosynthesis strategies
Leaf senescence
Osmotic
Ion homeostasis: adjustment:
Na+ extrusion, Accumulation of
Na+ exclusion, solutes
Na+ compartmentation
Adapted from Horie, T., Karahara, I. and Katsuhara, M. (2012). Salinity tolerance mechanisms in glycophytes: An overview with the central focus on rice plants. Rice. 5: 11; see also Munns, R. and Tester, M. (2008). Mechanisms of salinity
tolerance. Annu. Rev. Plant Biol. 59: 651-681 and Shabala, S. and Pottosin, I. (2014). Regulation of potassium transport in plants under hostile conditions: implications for abiotic and biotic stress tolerance. Physiol. Plant. 151: 257-279.
 General sodium tolerance strategy:
Keep sodium out of cytosol & shoot
OUT
1. Keep Na+ from
Na+ 4. Extrude Na+
IN Na+
via salt glands
entering plant / cells
“OUT”
K+ 5. Accumulate K+
2. Pump out any Na+ to maintain a high
Na+ that leaks in ratio of K+ to Na+

3. Compartmentation Na+ Compatible

of Na+ in vacuole solutes

6. Synthesize
compatible solutes for
osmotic balance 7. Prevent Na+
from moving into
the shoot and
leaves
 Ion pumps, channels & carriers
contribute to Na+ tolerance

PM-H+-ATPase Na+
ATP ADP + Pi
H+
SOS1, NHX8 H+
SOS1
H+ H+
Na+ ATP ADP+ Pi

H+ H+
PP 2 x Pi
HKT Na+

NSCC Na+ Na+

NHX H+-PPase VH+-ATPase

See Maathuis, F.J.M. (2014). Sodium in plants: perception, signalling, and regulation of sodium fluxes. J. Exp. Bot. 65: 849-858.
 The intersection of potassium
nutrition and sodium toxicity
K+ uptake

K+ext Na+ext

Na+ uptake

Na+ and K+ interfere with

each other’s uptake

When barley plants are grown

on 200 mM NaCl, they
accumulate Na+ at the
expense of K+ in their leaves

Cuin, T.A., Miller, A.J., Laurie, S.A. and Leigh, R.A. (2003). Potassium activities in cell compartments of salt‐grown barley leaves. J. Exp. Bot. 54: 657-661 with permission from Oxford University Press.
 Nitrogen: Unsur mineral paling
melimpah dalam tumbuhan
• Unsur paling melimpah di atmosfer
bumi
• Unsur yang paling banyak ke-4
dalam tanaman (setelah C, H dan
O).
• Seringkali sebagai mineralpembatas
untuk pertumbuhan tanaman

N terdapat dalam asam

amino (protein), asam
nukleat (DNA, RNA),
klorofil, dan molekul
kecil yang tak terhitung
jumlahnya
Nitrogen adalah salah
satu dari tiga
makronutrien utama
yang ditemukan pada
Blank, L.M. (2012). The cell and P: From cellular function to biotechnological application. Curr. Opin. Biotech. 23: 846 – 851.From: Buchanan, sebagian besar pupuk
B.B., Gruissem, W. and Jones, R.L. (2000) Biochemistry and Molecular Biology of Plants. American Society of Plant Physiologists.
 Nitrogen dapat ditemukan dalam
berbagai bentuk anorganik
Species Name Oxidation
State
R-NH2 Organic nitrogen, urea -3 NO2-
Nitrification
NH3, NH4+ Ammonia, -3
ammonium ion NO3-
Nitrate Aerobic
N2 Nitrogen 0 reduction reactions
NH3
N2O Nitrous oxide +1 Anaerobic
reactions
NO Nitric oxide +2 NO2- NO Nitrogen
N2O fixation
HNO2, NO2- Nitrous acid, +3
nitrite ion N2
NO2 Nitrogen dioxide +4
HNO3, NO3- Nitric acid, nitrate ion +5

Adapted from Robertson, G.P. and Vitousek, P.M. (2009). Nitrogen in agriculture: Balancing the cost of an essential resource. Annu. Rev. Environ. Res. 34: 97-125.
 Tumbuhan berperan penting dalam
siklus nitrogen global
Atmospheric pool of N2
Biological Industrial Atmospheric
fixation fixation fixation
5 Tg N / yr
Biological
NO3- fixation
(oceans)

denitrifying bacteria
Denitrification by
manure decomposition
NH4+
R-NH2 NO3-
NO3- Assimilation
NH4+
by plants
140
120 Tg N / yr 120 Tg N / yr
(50% Tg N / yr NH4+ NO2- NO3-
agricultural)
Nitrification by nitrifying bacteria

Adapted from Fowler, D., et al. (2013). The global nitrogen cycle in the twenty-first century. Phil. Trans. Roy. Soc. B: 368: 20130164
 Metabolisme nitrogen: penyerapan,
asimilasi dan remobilisasi
Remobilisasi Siklus karbon
Daur ulang asam Assimilasi TCA
Uptake amino,
Glutamate 2-oxoglutarate
fotorespirasi
NH4+ Glutamine-2-
oxoglutarate
NH4+ aminotransferase
Nitrite (GOGAT)
reductase
Glutamate
NO2 -
Nitrate
reductase Glutamine Glutamine
synthetase Penggabungan
NO3 - NO3 - (GS)
ke dalam asam
amino dan
R-NH2 senyawa yang
NH4+ mengandung
Assimilasi nitrogen lainnya
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.
 Sebagian besar tumbuhan meyerap
nitrogen dalam bentuk nitrat NO3-
Banyak prokariota mengoksidasi Tumbuhan merekduksi NO3- untuk
NH4+, sehingga kadar NH4 di tanah asimilasi menjadi senyawa organik
sering rendah
Energy Energy Energy Energy
released released consumed consumed

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

Nitrate Nitrite
Nitrification by nitrifying prokaryotes reductase reductase

Preferensi tanaman untuk NH4+ vs NO3- bervariasi

berdasarkan spesies, proses metabolisme lainnya, suhu,
air, pH tanah dll….

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.
 Tanaman memiliki transporter khusus
untuk NO3-, NH4 + dan bentuk N lainnya

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
Nitrat reduktase adalah enzim dengan
skema katalitik yang kompleks
NADH NAD+ Nitrat reduktase mereduksi
nitrat menjadi nitrit dengan
NO3- NO2- NADH bertindak sebagai
donor elektron

NO3-
NADH
Elektron berpindah dari
NADH ke FAD pada heme
ke kofaktor molibdenum
(Moco) mengikat 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.
 Nitrogen Assimilation I
• NO3- dapat diasimilasikan baik akar atau tunas,

– NO3- diserap oleh akar, disimpan dalam vakuola,

diasimilasi atau diangkut ke daun,
– NO3- direduksi menjadi nitrit,
– NO2- direduksi untuk membentuk amonium,
• menggunakan reaksi terang
fotosintesis untuk mengurangi
reaksi di daun, pengurangan
ferredoxin oleh metabolisme
plastid di akar,
• Penurunan ferredoxin oleh
metabolisme plastid di akar,
• NH4+ NH4 (dari matriks tanah, atau dari reduksi NO3-)
dimasukkan ke dalam asam amino untuk digunakan,
atau untuk transport N.
 Nitrate Toxicity
• NO3- → umumnya tidak beracun untuk
tanaman dan dapat dengan mudah
disimpan dan diangkut,
• Beracun dalam konsentrasi tinggi →untuk
organisme lain
– Methemoglobinemia (“blue baby
syndrome”),
• hati mereduksi nitrat menjadi nitrit
(atau pH yang lebih tinggi, melalui
bakteri dalam usus),
• nitrit mengoksidasi besi dalam
hemoglobin, (Fe2+ ---> Fe3+),
• menjadikan hemoglobin tidak dapat
mengikat O2,
– Nitrosamines,
• R2NNO or RNHNO,
• Sangat mutagenik dan
 Nitrogen Fixation Requires
Energy

Ammonia
and H2
Output.

8 reduced
ferredoxins
N2
8 protons

16 ATPs for hydrolysis

…why would bacteria associate with plants
Simbiosis Tumbuhan pada
asimilasi N2
– Alder (trees), Ceanothus (shrub),

• Frankia (bacteria),

– Sugarcane,

• Nostoc (bacteria),

– Azolla/Fairy Fern (water fern),

Ceanothus cordulatus; Mountain Whitethorn Frankia (bacteria
• Anaebena, (cyanobacteria)

Azolla Anabaena
Azolla/Anabaena/Rice/Humans
1. Spread Azolla, let
grow,
2. Break clumps,
drain paddy,
3. Stomp (or
machine smash),

4. Fixed nitrogen is
released to soil.
 shu (soybean)
Chou scholars (~1000 B.C.)
Soy Root Nodules: result from an
infection by Rhizobium.
Nitrogen Fixation
Rhizobium symbionts
 Rhizobium Infection I

if compatible?

1. Emerging root hair sends 3. Root hair grows and curls around the
chemical attractants (elicitors), bacterial colony,

2. Bacteria respond with a 4. Bacteria proliferate within the curl.

recognition signal,
 Sweet Talk?
3. Flavanoid binds 4. nod genes are expressed,
transcription factor (Nod D),
complex bids DNA,

5. Nod factor is produced

(a molecule with host
2. Bacterial receptor
specificity),
recognizes signal, and
transports it across cell
membrane,
6. Nod factor is recognized
by the host, in turn
1. Plant secretes a specific activating host genes for
elicitor (flavonoid). proper response.
 Rhizobium Infection II

5. Plant cell wall is degraded, 7. Infection thread reaches root hair

plasma membrane, fuses,
6. Plant plasma membrane
invaginates root hair cell, 8. Bacteria enter the apoplast.
 Rhizobium Infection III
9. New infection threads form,
10. Threads form toward “target
cells”,
11. Bacteria “bleb” off of the infection
thread, into the cytosol,
12. Bacteria are surrounded by a plant
membrane.
Nodule Formation
12. Bacteria are surrounded by a plant
Benang Bakteri
Rhizobium
Sel-sel
Yang sedang
membrane.
infeksi
membelah di
Korteks akar
- bacterial induce plant cell
Rambut akar
Bacteroid Sel-sel yang sedang
membelah pada
division in infected and
Yang terinfeksi 1
perisikel surrounding cells,
12
- cells in the pericycle begin
dividing, (similar to lateral root
formation).
13. Affected pericycle and
cortical derived cells
Nodul akar yang
Sedang berkembang
continue to divide until
the regions fuse.
Bacteroid
13

14
14. Vasculature forms between the
nodule and the plant stele,

- nitrogenous compounds are

carried to the plant,
Jaringan - nutrients to the bacteria.
Vaskular
Bacteroid nodul
Nitrogen Assimilation II

• Symbiosome,
– plant membrane, surrounding…

– one or more bacteria,

• once inside a symbiosome,
bacteria differentiate into
bacteroids,
• bacteroids may differentiate,
• Host Cell Synthesizes,
– transport proteins for the symbiosome
membrane,

– leghemoglobin, an oxygen binding

molecule,

– N assimilation enzymes.
 Bakteroid
Nodul Di dalam
vesikel

Akar 5 m

(a) Akar tumbuhan ercis (b) Bakteroid pada nodul

Akar kedelai
 Nitrogen Assimilation I vs. II

• Keuntungan simbiosis bagi tumbuhan.

– kecuali terjadi kekurangan nitrogen,
tumbuhan tanpa infeksi rhizobium pada
umumnya lebih baik dari tumbuhan yang
terinfeksi,
– Secara umum,
• high NO32-, no R (best growth),

• high NO32-, R (good growth),

• low NO32-, R (good growth),

• low NO32-, no R (poor growth).

I II