Materi Belajar
Materi Belajar
Materi Belajar
Soils that contain a harmful amount of salt are often referred to as salty or saline soils. Salt in seawater is merely dissolved in the water, not chemically bonded to it.
When water evaporates (one molecule at a time), only pure water returns to the atmosphere. Salt and other impurities are left behind.
Salinisasi merupakan proses terjadi peningkatan garam mudah larut (NaCl, Na2CO3, Na2SO4) yang tinggi dalam tanah, sehingga berpengaruh terhadap pertumbuhan
dan perkembangan tanaman.
Though the material that gives seawater its salty flavour is composed of many substances, sodium chloride, or common salt, is by far the predominant compound.
Sodium Chloride dissolves in water into its ions, which are present at much higher levels than components of any other salt. Sodium and Chloride together make up
around 85 per cent of all dissolved ions in the ocean. Ocean salt primarily comes from rocks on land and openings in the seafloor. Salt in the ocean comes from two
sources: runoff from the land and openings in the seafloor. Rocks on land are the major source of salts dissolved in seawater.
It takes just slightly more energy to separate the ions from one another than is released from the water molecules surrounding the ions. This
means just slightly more energy must be put into the solution than is released back into the solution; therefore, dissolving table salt in water is
endothermic
The main difference between exothermic and endothermic reactions is that an endothermic reaction absorbs energy in the form of heat from its
surroundings, whereas an exothermic reaction releases energy to the surroundings. In endothermic dissolution reactions, the net energy from
breaking and forming bonds results in heat energy being absorbed into the system as the solute dissolves.
Salt in the ocean comes from two sources: runoff from the land and openings in the seafloor. Rocks on land are the major source of salts dissolved
in seawater. Rainwater that falls on land is slightly acidic, so it erodes rocks. This releases ions that are carried away to streams and rivers that
eventually feed into the ocean. Many of the dissolved ions are used by organisms in the ocean and are removed from the water. Others are not
removed, so their concentrations increase over time
Irrespective of the source of the seawater, salt obtained by the evaporation of seawater has the following composition: sodium chloride 77.76
percent, magnesium chloride 10.88 percent, magnesium sulfate 4.74 percent, calcium sulfate 3.60 percent, potassium chloride 2.46 percent, magnesium bromide
0.22 percent, and calcium carbonate 0.34 percent.
Seawater is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of ~3.5%. This means that for every 1 litre (1000 mL) of seawater
there are 35 grams of salts (mostly, but not entirely, sodium chloride) dissolved in it.
Salinitas tidak hanya menurunkan produksi pertanian, tetapi juga mempengaruhi sifat fisikokimia dan keseimbangan ekologi pada tanah. Akibatnya, salinitas akan
memberikan dampak buruk bagi hampir semua aspek budi daya tanaman, seperti perkecambahan, pertumbuhan vegetatif, perkembangan, dan reproduksi (Aderoju
et al., 2013).
Evapotranspiration
is the name given to
the total water loss
to the atmosphere
from a land surface,
usually expressed in
units of depth; it
includes the water
vapor evaporating
from the soil surface
and from the liquid
water on plant
surfaces together
with that transpired
from within plant
surfaces.
Alluvial soil: Alluvial soil are formed by the deposition of divers which can be mostly found in Northern plain regions and the regions that surrounded by
Narmada and Tapi rivers and also in some parts of Northern Gujarat. Black Soil: Black soil is formed of by the action of volcanic rock and lava-flow.
The diversity of alluvial soils results in a complex array of potential soil classifications. Recent alluvial soils that lack significant development of surface or
subsurface diagnostic features are classified into the order of Entisols (US Department of Agriculture Soil Taxonomy System) or Fluvisols (Food and
Agriculture Organization World Reference Base for Soil Resources).
In the Soil Taxonomy System, the formative element ‘fluv’ is used to connote the alluvial origin and stratified nature of recent alluvial soils. Better-drained
Entisols that have high organic carbon levels (more than 0.2%) at depth (125 cm) or an irregular decrease in organic carbon with depth (see Figure 3) are in
the suborder of Fluvents. Fluvents are further subdivided into ‘great groups’ by soil moisture regimes (e.g., Udifluvents, Torrifluvents, etc.). Alluvial Entisols
that are saturated for prolonged periods during a normal year are Fluvaquents. In arid climates, the organic carbon level in the soil is often low; thus, many
youthful alluvial soils in arid climates are classified as Torriorthents.
With greater landform stability and soil development, alluvial soils can develop into a myriad of different soils and thus can occur in all of the other soil
orders of the Soil Taxonomy System. Alluvial soils with mollic epipedons and high base status throughout are Mollisols. Alluvial soils in arid climates that
develop subsurface diagnostic horizons (e.g., argillic, calcic, gypsic) are Aridisols. Other alluvial soils with ochric or umbric epipedons
are Inceptisols, Alfisols, Ultisols, Oxisols, and Spodosols, depending on the diagnostic subsurface horizons and properties present and the degree of leaching.
Alluvial soils subject to cryoturbation are Gelisols; alluvial soils that accumulate deep organic soil materials are Histosols. Vertisols and Andisols are also
possible, depending on the mineralogical composition, texture, and the degree of soil development.
Alfisols (aluminium iron soil) 2. Andisols 3. Aridisols (dry soil) 4. Entisols (recent soil)
Alfisols merupakan tanah yang kaya akan Andisols merupakan tanah yang berasal dari Aridisols merupakan tanah kering yang Entisols merupakan tanah baru, yaitu tanah
bahan organik, besi, air alumunium, dan endapan abu vulkanik gunung api. terbentuk di gurun dan semi gurun (di daerah yang sangat muda dan banyak Gelisols
struktur tanahnya berlapis-lapis. Merupakan Penyebaran tanah ini hanya terdapat di beriklim kering atau daerah bayangan hujan) ditemukan pada daerah beriklim sangat
tanah yang berkembang pada daerah hutan sekitar lereng-lereng gunung berapi aktif. Ciri- dengan struktur tanah berlapis-lapis. Ciri-ciri kering atau daerah beriklim dingin. Tanah ini
dan sabana. Ciri-ciri tanah alfisols adalah: ciri tanah andisols adalah : tanah aridisols adalah: belum atau sedikit mengalami perkembangan
warnanya abu-abu berwarna coklat bercampur hitam berwarna kemerah-merahan horison sehingga tidak berlapis. Ciri-ciri tanah
mengandung lapisan tanah liat atau mengandung banyak mineral tanah terdapat lapisan kapur di bawah entisols adalah:
lempung sangat baik bagi kegiatan pertanian permukaannya terdapat di dataran banjir dari
tanahnya subur dan sangat baik kandungan air dan bahan organik sungai-sungai dan bukit-bukit pasir
untuk kegiatan pertanian sedikit kandungan bahan organik sedikit
vegetasi di tanah tersebut sedikit tanahnya subur.
digunakan untuk padang rumput.
5. Gelisols 6. Histosols (organic soil) 7. Inceptisols (beginning soil) 8. Mollisols (Soft Soil)
Gelisols merupakan tanah dengan ciri-ciri Histosols adalah tanah yang mengandung Inceptisols merupakan tanah muda, yaitu Mollisols adalah tanah halus, yaitu tanah
sebagai berikut : bahan tumbuh-tumbuhan yang sudah tanah yang agak lebih tua dari entisols. Tanah halus kering, berwarna coklat, merah, dan
lapisan top soil merupakan bahan membusuk dan air. Tanah ini bersifat jenuh ini mulai mengalami perkembangan pada tiap hitam. Tanah ini berkembang pada daerah
organik air dan menciptakan kondisi anaerob dan horisonnya sehingga perbedaan tiap lapisan padang rumput. Tops soilnya berwarna gelap
lapisan subsoil mengalami menyebabkan terjadinya akumulasi bahan mulai terlihat jelas. Ciri-ciri tanah inceptisols dan kaya akan humus. Ciri-ciri tanah mollisols
pembekuan organik. Ciri-ciri tanah histosols adalah: adalah:
banyak ditemukan di daerah lintang adalah: terdapat di daerah tundra atau mengandung banyak bahan organik
tinggi atau di wilayah tundra terdapat di bekas danau, rawa, dingin dan air
daerah berpaya yang selalu tanah tersebut asam dan banyak struktur tanahnya berlapis
dipenuhi tumbuh-tumbuhan dan mengandung bahan organik terdapat di daerah semi arida
lumpur digunakan untuk pertanian atau tanahnya paling subur, baik untuk
tanahnya subur tanaman rumput untuk ternak pertanian
dapat dijadikan tanah pertanian baik untuk pertumbuhan rumput
yang baik bila dilengkapi dengan ternak.
pengairan atau drainase yang baik
di Indonesia terdapat di pantai
Kalimantan dan pantai timur
Sumatera.
Tanah rawa (3,5-5 pH)
9. Oxisols 10. Spodosols (Ashy Soil) 11. Ultisols / last (Ultimate Soil) 12. Vertisols (Turned Soil)
Oxisols merupakan tanah purbakala, yaitu Spodosols adalah tanah tua, yaitu tanah yang Ultisols, yaitu tanah yang sudah mengalami Vertisols yaitu tanah yang menjadi matang,
tanah yang mengandung banyak oksigen, berlapis, mengandung silika, aluminium, besi, perkembangan masa tua (terakhir). Tanah ini dengan lapisan yang tebal (dalam). Ciri-ciri
banyak tanah liat, tetapi bahan organiknya dan bahan organik. Ciri-ciri tanah spodosols mengalami pencucian di daerah yang basah tanah vertisols adalah:
sedikit. Ciri-ciri tanah oxisols adalah: dan hangat. Ciri-ciri tanah ultisols adalah: mengandung tanah liat, kapur, dan
adalah berwarna abu-abu atau kemerahan terdiri dari lapisan tanah liat, bahan organik
warnanya kemerah-merahan terdapat di daerah beriklim sejuk alumunium, dan bahan organik terdapat di daerah beriklim panas
tidak cocok untuk pertanian hingga dingin terdapat di daerah beriklim tropik dengan musim basah dan kering
terdapat di daerah iklim panas dan di daerah humida, misal di hutan yang lembab baik untuk pertanian apalagi kalau
tropik dengan curah hujan yang konifer dengan tingkat keasaman terbentuk pada daerah dengan dipupuk.
banyak tinggi batuan induk yang sudah tua Tanah kapur pH >8
mineral tanah didominasi oleh besi tanahnya bersifat asam sehingga tidak baik untuk pertanian, kecuali
dan aluminium kurang mampu menahan air kalau dipupuk
cenderung tidak subur dan tidak
cocok untuk pertanian.
Fertile riverine alluvial soil and the clayey loam texture of the new alluvial soil is the most suitable for rice cultivation
Rice is typically grown in bunded fields that are continuously flooded up to 7−10 days before harvest. Continuous flooding helps ensure sufficient water and
control weeds. Lowland rice requires a lot of water. On average, it takes 1,432 liters of water to produce 1 kg of rice in an irrigated lowland production system.
Total seasonal water input to rice fields varies from as little as 400 mm in heavy clay soils with shallow groundwater tables to more than 2000 mm in coarse-
textured (sandy or loamy) soils with deep groundwater tables.
Kenapa tanah gambut pH nya rendah? Tanah bereaksi masam (pH rendah 3-5) adalah karena tanah kekurangan Kalsium (CaO) dan Magnesium (MgO), ini
disebabkan oleh: Curah hujan tinggi, pada daerah dengan iklim tropika basah, dengan curah hujan yang tinggi, secara alami tanah akan menjadi masam akibat
pencucian unsur hara yang ada. Tingkat kemasaman tanah gambut berhubungan erat dengan kandungan asam-asam organik, yaitu asam humat dan asam fulvat
(Andriesse, 1974; Miller dan Donahue, 1990).
Berdasarkan Peta Tanah Eksplorasi Indonesia (Puslitbangtanak 2000), Ordo tanah yang dijumpai terdiri dari 10 Ordo, dan tidak dijumpai ordo tanah Aridisol, hal
tersebut dapat dimengerti karena di Indonesia tidak dijumpai adanya rejim kelembaban tanah aridic yang merupakan salah satu penciri utama dari Ordo tanah
Aridisol (Gambar 3).
Soil dispersion causes clay particles to plug soil pores,
resulting in reduced soil permeability. When soil is
repeatedly wetted and dried and clay dispersion occurs, it
then reforms and solidifies into almost cement-like soil
with little or no structure. The three main problems caused
by sodium-induced dispersion are reduced infiltration,
reduced hydraulic conductivity, and surface crusting. Salts
that contribute to salinity, such as calcium and magnesium,
do not have this effect because they are smaller and tend
to cluster closer to clay particles Calcium and magnesium
will generally keep soil flocculated because they compete
for the same spaces as sodium to bind to clay particles.
Increased amounts of calcium and magnesium can reduce
the amount of sodium-induced dispersion.
Menurut Strawn et al. (2015) tanah salin adalah tanah yang banyak mengandung garam dan dicirikan oleh nilai Electrical Conductivity (EC) > 2 dS/m atau lebih
dalam larutan tanah.
Sementara itu Van Wambeke dan Forbes (1986) mengemukakan bahwa tanah salin atau tanah bergaram dicirikan oleh nilai persentase natrium dapat ditukar
(ESP, exchangable sodium precentase) > 15% atau nilai bandingan adsorpsi natrium (SAR, sodium adsorption ratio) > 13.
Pada lahan salin kadar NaCl berkisar antara 2-6 %. Kandungan Na yang sangat tinggi di dalam tanah akan berakibat buruk bagi sifat fisika tanah karena akan
menyebabkan pelarutan liat (clay dispersion) yang lebih jauh lagi dapat mengakibatkan penyumbatan dan pembentukan kerak pada kesarangan tanah sehingga
kepadatan tanah meningkat. (The forces that bind clay particles together are disrupted when too many large sodium ions come between them. When this
separation occurs, the clay particles expand, causing swelling and soil dispersion)
Apabila semua kapasitas adsorpsi tanah telah dijenuhi oleh ion Na+ , akan terjadi fenomena “Tanah Larut” (dispersive soils). Penjenuhan kapasitas adsorpsi
menyebabkan lempeng-lempeng dalam partikel liat saling tolak-menolak sehingga melarut (disperse) dalam air dalam bentuk koloidal berukuran submikron atau
ångström.
Penurunan laju fotosintesis juga dapat dikaitkan dengan prilaku stomata. Pada tanaman yang mengalami stress garam, dimana juga mengalami defisiensi air,
kosentrasi CO2 pada kloroplas menurun karena berkurangnya konduktansi stomata. Penurunan aktivitas fotosintesis tanaman tersebut akan mempengaruhi
pembentukan berat kering sehingga menganggu pertumbuhan tanaman.
Kebanyakan tanaman yang menderita stress garam menunjukkan penurunan pertumbuhan dan juga hasil tanaman. Pertumbuhan dan hasil tanaman budidaya
umumnya mengalami penurunan pada EC tanah 4 dS/m atau lebih, bahkan tanaman yang sensitif dapat terpengaruh pada EC 3 dS/m.
Tanda-tanda tanaman yang terkena stress garam antara lain menjadi kerdil, kesehatan tanaman terganggu, warna tanaman berubah dan hasil tanaman menurun
(McWilliams 2003). Selanjutnya Sukarman et al. (1998) mengemukakan bahwa pada tanah salin sebagian besar tanaman budidaya tidak bisa hidup dengan baik.
Hal tersebut disebabkan karena tekanan osmotik tanah sangat tinggi dan tingginya kandungan anion khlorida (Cl- ) yang bersifat toksik bagi tanaman.
Salinitas yang tinggi menyebabkan pertumbuhan tanaman terhambat karena turunnya tekanan osmotik, sehingga menyulitkan pengambilan unsur hara oleh
akar.
Sodisitas tinggi menyebabkan keracunan Na dan ion-ion sejenis, seperti Boron dan Molibdenum. Disamping itu, terdapat efek tidak langsung dari keduanya
berupa peningkatan nilai pH tanah yang menyebabkan imobilitas beberapa unsur hara penting seperti Ca, Mg, P, Fe, Mn, dan Zn sehingga unsurunsur tersebut
tidak dapat di ambil oleh akar tanah
Ada dua cara menentukan salinitas air yaitu dengan menentukan Total Disolved Salt (TDS) dan Electric Conductivity (EC).
Siemens (simbol: S) adalah satuan turunan dari konduktansi listrik, suseptansi listrik, dan masuk listrik dalam Satuan Sistem Internasional (SI).
Hasil pengukuran menggunakan EC dinyatakan dalam μS/cm (mikroseimen/cm) dan dapat dikonversikan ke mg/l dengan menggunakan grafik yang disajikan oleh
Hansen et al. (1992).
Kekeringan akibat perubahan iklim juga menjadi salah satu penyebab terjadinya proses salinisasi. Hasil riset menunjukkan bahwa ada hubungan antara
pemanasan global dengan peningkatan salinitas lahan karena pemanasan global memacu lelehnya es di kutub bumi sehingga permukaan air laut naik, akibatnya
dataran yang lebih rendah akan tenggelam dan tekanan air laut ke daratan juga meningkat (Dailidienė dan Davulien, 2008).
Sembiring dan Gani (2007) menyatakan bahwa akibat peningkatan salinitas di lahan-lahan sawah sekitar Pantura, banyak petani yang mengalihfungsikan
lahannya ke usaha ladang garam atau perikanan tambak bahkan memberakan lahannya karena usahanya yang sudah tidak menguntungkan lagi
Mikroiza vs pgpr
- Mikoriza lekat sm akar trs miceliumnya menyerap air
- •The majority of these studies were conducted in controlled growth chamber or
greenhouse conditions, while very few have been conducted in field conditions. We need more
data regarding the variability or stability of AMF-conferred salinity tolerance before advocating
for farm-level applications.
Humic acid increases nutrient uptake, drought tolerance, and seed germination. It increases the microbial activity in the soil, making it an excellent root
stimulator. Humic acid increases the availability of nutrients in our fertilizers and in those already existing in your soil. The HAs are built of structures with both
hydrophobic and hydrophilic properties which determine, e.g., the solubility and the susceptibility of HAs to biodegradation and affect their sorption potential
(Wen et al. 2007).
Humic substances are recognized as the most important component of soil as they influence the soil structure and play the main role in the chelation of mineral
elements. The structure, which shows important effects on the physical properties of soil, is often measured by the stability of soil aggregates (Six et al. 2000;
Bronick and Lal 2005). Soil aggregate stability is a crucial soil property affecting soil sustainability (Amézketa 1999). The size distribution and stability of soil
aggregates positively correlates with the SOM and clay minerals (Tisdall and Oades 1982; Six et al. 2004). There was also found a positive relationship between
the stability of soil aggregates, cation exchange capacity (CEC), and the content of humic substances
Since these organic fractions are from diverse organic materials they presumably have a different structural composition, and this might be the reason for their
different behaviour. The greater degree of aliphaticity and the higher content in free carboxyl groups of molecules forming the FHM fraction would lead to the
formation of more numerous linkages. As a result of this a mineral-organic material might be formed, which would allow the infilling of planes and the binding
between large particles to take place, thus giving rise to more-stable and larger aggregates. However, as FHP consists of more-polymerized molecules that have a
smaller proportion of free carboxyl groups, the formation of mineral-organic material would not happen so quickly when this fraction is used and, in addition,
only binding between small particles or clay would occur thereby increasing the formation of smaller aggregates. The heterogeneous distribution of organic
material in the aggregate would cause the increase in vughs. These processes are also manifest when comparing soils, and so the higher clay content in the clay
soil would allow this soil to have a greater binding capacity which might favour the aggregate formation.
- Ada 4 langkah utama dalam proses biosintesis EPS: 1) penyerapan, aktivasi dan konversi monosakarida menjadi
nukleotida gula di sitoplasma; 2) metabolisme, perakitan unit berulang nukleotida gula dengan penambahan berurutan
setiap unit pada molekul pembawa lipid dengan bantuan glikosiltransferase; 3) polimerisasi unit berulang yang terjadi
di sisi periplasma membran plasma menjadi bentuk polisakarida, dan 4) eksudasi EPS ke permukaan sel
- Komposisi EPS Pseudomonas aeruginosa di bawah tekanan garam umumnya didominasi oleh rhamnosa, manosa, dan
trehalosa. Manosa, rhamnosa, dan trehalosa dapat membantu meningkatkan toleransi terhadap stres garam dengan
menyediakan sumber karbon, meningkatkan 20 retensi air. Trehalosa juga berperan dalam melindungi enzim seluler
dengan menstabilkan membran sel selama kondisi stress (Tewari & Arora, 2014)
Soil water retention capacity is the maximumamount/quantity of water
that a soil can hold or retain. Itis very important property with respect to
formers point ofview as well as plant growth. If a soil can hold largeamount
water it decrease the irrigation frequency of cropand also plant grow well
this type of soil. It wasinvestigated that biochar application boast up
theavailable water content of the soil up to 97 percent andsaturated water
contents 56 percent (Uzoma et al.,2011). Laird et al., (2010) described that
the biocharamended soil retained 15 % more moisture contents
ascompared controlled treatment. According to E. H. Tryon(1948) biochar
application into the soil increased thewater retention capacity of the soil
but it is texturedependent. There is significantly increased in soil
waterretention capacity in case of sandy soil by biocharapplication but
there little and no increased in loamy andclay soil respectively. Herth et al.,
(2013) describedexperimentally that biochar application increased
thewater retention capacity of the soil because it increasesoil porosity and
also due to adsorptive nature of biochar.Uzoma et al., (2011) said that
there were hydrophilicfunctional groups present on the surface of the
graphynesheet of the biochar and also on the pores.
Tryptophan is naturally secreted in root exudates of tomato plants and most of the auxin found in the rhizosphere is believed to
come from the biosynthesis by microorganisms (Kamilova et al. 2006). Antagonistic Pseudomonas sp. PsM6 and PjM15 and
growth-promoting strains belonging to the same genera utilized tryptophan (Trp) as a precursor for IAA production (Patten and
Glick 1996; Rajkumar and Freitas 2008) and growth (Rajkumar and Freitas 2008). For rhizobacteria from wheat, rice and maize
(Azospirillum lipoferum, Acinetobacter genospecies, Bukholderia vietnamiensis, Enterobacter cloacae, Pseudomonas
aeruginosa and Pseudomonas putida), IAA biosynthesis may occur in the presence as well as in the absence of tryptophan
(Asghar et al. 2002; Khalid et al. 2004). Bacterial IAA is generally synthesized via two pathways, the Trp-independent and the
Trp-dependent by sequential conversion of tryptophan to various intermediates.
Aseptic tomato and radish roots were found to exude, respectively, 2.8–5.3 and 290–390 ng tryptophan per seedling per day.
The inoculation of radish plants with rhizosphere pseudomonads increased the root biomass by 1.4 times.
Ethylene is a simple gaseous hormone which plays multiple roles in regulation of plant growth and development, and also serves as a key modulator between
plant response to environmental stresses and normal growth (Abeles et al., 1992). Under salinity and some other stresses, ethylene production is quickly
stimulated (Morgan and Drew, 1997). In fact, several key steps of ethylene biosynthesis could be affected by salinity and other stresses. Ethylene in plant is
synthesized through three enzymatic reaction steps: methionine is converted to S-adenosyl-methionine (S-AdoMet) by S-AdoMet synthetase; then the direct
precursor of ethylene ACC is synthesized from S-AdoMet by ACS (ACC synthase); and finally ethylene is produced through the oxidation of ACC by ACO (ACC
oxidase) (reviewed in Lin et al., 2009). S-AdoMet is also the precursor for the synthesis of polyamines, which also plays a role in plant response to biotic and
abiotic stresses. The by-product MTA (5′-methylthioadenosine) from the second step is recycled to methionine through the Yang Cycle to maintain a stable
methionine pool even when ethylene is rapidly synthesized (Yang and Hoffman, 1984; Kende, 1993).
As a rate-limiting enzyme, ACS is the major target for regulation of ethylene production under stresses (Yang and Hoffman, 1984; Kende, 1993; Bleecker and
Kende, 2000; Wang et al., 2002). There are eight functional ACS genes in Arabidopsis. They have redundant role in ACC biosynthesis (Tsuchisaka and Theologis,
2004; Tsuchisaka et al., 2009), while each with unique function in the regulation of plant growth and development. The regulation of ACSs under stress occurs at
both transcriptional and post transcriptional levels. Under salinity, transcripts of ACS2 and ACS7 in Arabidopsis were increased dramatically (Achard et al., 2006).
Through GUS staining, Arabidopsis ACS5 and ACS7 promoters were found to be induced by salinity (Wang et al., 2005). A more systematic research revealed that
the MAPK cascade-induced by stress might activate WRKY33 which then promoted the expression of ACS2/ACS6 genes in Arabidopsis (Li et al., 2012). In
tobacco, the transcripts of ACS1 were induced by salinity (Cao et al., 2006). In cotton, a series of ACSs were up-regulated under both short- and long-time salinity
condition (Peng et al., 2014b). Recent work in Arabidopsis found that four ACSs (ACS2, ACS6, ACS7, and ACS8) were induced by high salinity, while a moderate
low salinity pretreatment (known as salt acclimation) alleviated this induction (Shen et al., 2014). This result provides us a cue that promotion of ethylene
production by a strong and sudden salinity stress might have multiple effects for plant response to salinity. It seems that no extra ethylene was needed for the
process of salt acclimation, because there were no changes on the expression of genes related to ethylene biosynthesis (Shen et al., 2014). Four ACSs were up-
regulated no matter under NASS (non-acclimated salt stress) or SASS (salt acclimated salt stress; Shen et al., 2014), indicating that promotion of ethylene
production is still necessary for plant adaption to the stress condition. Nevertheless, transcripts of ACSs under SASS were less than those under NASS ( Shen et
al., 2014). These results suggest that a tight control of ethylene biosynthesis might be important for plant response and adaption to salinity stress.
Besides transcriptional regulation, some ACSs are also regulated post-transcriptionally under salinity, mainly through stress-activated MAPK (mitogen-activated
protein kinase) cascades which phosphorylate ACSs protein and then prevent them from 26S proteasome-mediated degradation. Under biotic and abiotic
stresses, Arabidopsis MPK6 was activated rapidly to phosphorylate ACS2 and ACS6 to elevate ethylene production (Liu and Zhang, 2004). In tomato, it was
shown that calcium-dependent protein kinases (CDPKs)-mediated phosphorylation stabilized ACS2, leading to increased ACC content in stressed tissues ( Tatsuki
and Mori, 2001; Kamiyoshihara et al., 2010). In Arabidopsis, loss-of-function of MPK6 diminished the effects of salt acclimation, leading to more sensitivity to
high salinity (Shen et al., 2014).
Another key ethylene biosynthesis enzyme ACO is also regulated by salinity. Under high salinity, both ACC content and ACO activity were increased to enhance
ethylene release in Cicer arietinum root (Kukreja et al., 2005). In cotton, several ACOs were up-regulated under both short- and long-time salt treatment (Peng et
al., 2014b). From these results, we can propose that plants under salinity and other abiotic and biotic stresses tend to produce more ethylene mainly through
enhancing ACSs and ACOs. Nevertheless, transcripts of ACO1 in wheat were decreased under salinity and other abiotic stresses (Chen et al., 2014).
Although ACSs and ACOs are usually up-regulated under salinity, these key enzymes for ethylene biosynthesis may play negative roles in plant salinity-response.
In Arabidopsis, loss-of-function of ACS7 conferred less ethylene emission, promoted vegetative growth and enhanced tolerance to salinity (Dong et al., 2011).
Constitutive expression of wheat ACO1 in Arabidopsis led to salinity-sensitivity, possibly through increasing the expression of AtMYB15 while suppressing some
stress-responsive genes like AtRAB18, AtCBF1, and AtCBF3 (Chen et al., 2014). Expression of MKK9 in transgenic plants activated the endogenous MPK3 and
MPK6 kinases, promoted the synthesis of ethylene and camalexin, and finally conferred increased sensitivity to salinity, whereas loss-of-function
mutant mkk9 showed enhanced tolerance to salinity (Xu et al., 2008). Recent work in rice found that a lectin receptor-like kinase SIT1 (salt intolerance 1)
mediated salinity sensitivity through regulation of ethylene homeostasis (Li et al., 2014a). In presence of salinity stress, SIT1 was activated rapidly to
phosphorylate MPK3/6 and then promote ethylene production (Li et al., 2014a). Overexpression of SIT1 in rice and Arabidopsis both conferred increased
sensitivity to salinity, while loss-of-function of MPK3/6 alleviated this effect (Li et al., 2014a). Expression of SIT1 in rice and Arabidopsis also enhanced the ROS
accumulation in an MPK3/6- and ethylene signaling-dependent manner (Li et al., 2014a). All indicate a SIT1-MPK3/6 cascade mediates salt sensitivity by
regulating ethylene homeostasis. In addition, plant growth-promoting rhizobacteria (PGPR) produced AcdS (ACC deaminase), which facilitated the growth and
stress tolerance of hosts via a reduction in levels of ethylene (Ali et al., 2014; Barnawal et al., 2014; Kim et al., 2014). Transgenic Arabidopsis expressing AcdS
showed reduced sensitivity to exogenous ACC but increased tolerance to high salinity. In contrast, AcdS-silenced Trichoderma mutants were less effective in
promoting plant tolerance to salinity, indicating that Trichoderma can also ameliorated plant growth under salinity stress by decreasing the ethylene
biosynthesis in hosts (Brotman et al., 2013).
In contrast to the studies above, some other works showed that ethylene biosynthesis has positive effects on salinity-response. As stated above, MPK6, a key
regulator of ethylene biosynthesis under stresses, was necessary for salt acclimation (Shen et al., 2014). That means plants may need MPK6 to stabilize ACSs and
maintain a relative high ethylene level to accomplish the salt acclimation. The BTB ubiquitin ligases ETO1, EOL1, and EOL2 could interact with ACS5, promoted its
ubiquitination and then accelerated its degradation (Christians et al., 2009). The expression of ETOL1 in rice was induced under 200 mM NaCl treatment (Du et
al., 2014). In Arabidopsis, loss-of-function of ETO1 increased ethylene production and improved seedling tolerance to soil-salinity. Lack of ETO1 reduced root
Na+ influx and so restricted root-to-shoot delivery of Na +, and these effects were associated with increased RBOHF-dependent ROS accumulation in root stele. In
addition, loss-of-function of ETO1 enhanced the tissue K + status through an RBOHF-independent manner associated with increased transcripts of the K +-
transporter HAK5 (Jiang et al., 2013).
Iron is an essential element for the growth of almost all known organisms. It is required for electron transport
pathways (ferredoxins) as well as many metallo-enzymes (oxido-reductases, cytochromes and non-heme
oxidases) and in higher animals for oxygen transport (hemoglobin). The majority of iron in the environment is in
the form of insoluble ferric oxide/hydroxide complexes and within organisms iron is bound as a cofactor to
enzymes or stored as cluster complexes within proteins such as ferritin. As a result, many saprophytic and
pathogenic microrganisms have evolved methods of sequestering iron from their environment [1]. One of the
ways in which they achieve this is by producing small molecules with a high affinity for ferric iron known as
siderophores, whose biosynthesis is induced by intracellular iron deficiency and which are secreted into the
environment to scavenge iron [2]. The resulting iron complex is recognized by specific receptors on the microbial
cell surface and is actively transported into the cell [1,2]. Siderophores usually form octahedral (hexacoordinate)
complexes with ferric iron, employing a variety of func- tional groups as bidentate ligands including hydroxa-
mates, a-hydroxycarboxylates and catecholates, as well as polydentate phenolate/nitrogen heterocycle/carboxy-
late combinations. More than five hundred siderophores have been structurally characterized and there has
been growing interest in their biosynthesis because they often function as virulence factors in pathogens. Thus
enzymes involved in siderophore biosynthesis are potential targets for the development of antimicrobial
therapies [3]. Siderophores are broadly classified into two groups based on the kind of enzymatic machinery
involved in their biosynthesis. One group depends on non-ribosomal pep- tide synthetase (NRPS) modular
multienzymes [4], whereas the other is NRPS-independent [5]. Here we outline recent developments in the
elucidation of the biosynthetic pathways to these important natural products.
Phosphorus mineralization refers to the solubilization of organic phosphorus and the degradation of the
remaining portion of the molecule. One important theory proposed by Halvorson et al. (1990) for the
solubilisation of organic P is the sink theory. This refers to continuous removal of P that result in the
dissolution of Ca-P compounds. Consequently, the decomposition of P in organic substrates is consistently
correlated with the P content in the biomass of PSM (Dighton and Boddy, 1989). This biological process
plays an important role in phosphorus cycling. Different groups of enzymes are involved in this. The first
groups of enzymes are those that dephosphorylate the phosphor-ester or phosphoanhydride bond of
organic compounds. They are non-specific acid phosphatases (NSAPs). The most studied among these
NSAPs enzymes released by PSM, are the phosphomonoesterases also referred to as phosphatases
(Nannipieri et al., 2011). These enzymes can either be acid or alkaline phosphomonoesterases (Jorquera
et al., 2011). The pH of most soils where phosphate activities were reported ranges from acidic to neutral
values. This signifies that acid phosphatases play the major role in this process (Rodríguez and Fraga,
1999).
Another enzyme produced by PSM in the process of organic P mineralization is phytase. This enzyme is
responsible for the release of phosphorus from organic materials in soil (plant seeds and pollen) that are
stored in the form of phytate. Phytate degradation by phytase releases phosphorus in a form that is
available for plant use. Plants generally cannot acquire phosphorus directly from phytate, however, the
presence of PSM within the rhizosphere may compensate for a plant’s inability to otherwise acquire
phosphorus directly from phytate (Richardson and Simpson, 2011).
many experiments, the principal mechanism is the production of mineral dissolving compounds
such as organic acids, siderophores, protons, hydroxyl ions and CO2 (Rodríguez and Fraga,
1999; Sharma et al., 2013). Organic acids produced as described in Figure 2 together with their
carboxyl and hydroxyl ions chelate cations or reduce the pH to release P ( Seshachala and
Tallapragada, 2012); The organic acids are produced in the periplasmic space by the direct
oxidation pathway (Zhao et al., 2014). The excretion of these organic acids is accompanied by a
drop in pH that results in the acidification of the microbial cells and the surroundings, hence, P ions
are released by substitution of H + for Ca2+ (Goldstein, 1994). Surprisingly, Asea et al.
(1988) discovered that no correlation exists between the pH and the amount of P solubilized.
Hence Illmer and Schinner (1995) proposed the theory of acidification by H +. They explained that
H+ released is associated with cation assimilation. For example, assimilation of NH 4+ together with
H+ excretion brings about P solubilisation (Illmer and Schinner, 1995). An alternative mechanism to
organic acid production for solubilization of mineral phosphates is the release of H + to the outer
surface in exchange for cation uptake or with the help of H + translocation ATPase (Rodríguez and
Fraga, 1999). It was also reported that the assimilation of NH4 + within microbial cells is
accompanied by the release of protons and this results in the solubilization of phosphorus without
the production of any organic acids (Sharma et al., 2013). Of all the organic acids, gluconic acid is
the most frequent agent of mineral phosphate solubilization; it chelates the cations bound to
phosphate, thus making the phosphate available to plants. Gram-negative bacteria solubilize
mineral phosphate by direct oxidation of glucose to gluconic acid (Goldstein, 2000).
Pyrroloquinoline quinone (PQQ) acts as a redox cofactor in glucose dehydrogenases (GDH) resulting
in phosphate solubilisation (Rodríguez et al., 2000).
Natrium (Na+) juga penting untuk fiksasi karbon pada tanaman C4, seperti tanaman jagung, tebu dan sorgum. Sedangkan tanaman C3 diantaranya adalah
tanaman padi, gandum, kedelai. Pemasukan Na+ pada kondisi salin akan mengubah lintasan fotosintesis dari C3 menuju C4. Hal tersebut juga terjadi pada
tanaman jagung di mana Na + berpengaruh pada keseimbangan antara enzim fosfo enol piruvat karboksilase dan riboluse bifosfat karboksilase. Berlimpahnya
Na+ dan Cl- dapat mengakibatkan ketidakseimbangan ion sehingga aktivitas metabolisme dalam tumbuhan menjadi terganggu
Soil enzyme activities decreased with increasing ECe; however, the degree of inhibition varied among the enzymes assayed and the nature
and amounts of salts added. Dehydrogenase activity was severely inhibited by salinity, whereas, the hydrolases showed a much lesser
degree of inhibition. Generally, the inhibition of soil enzyme activities by the salt solutions decreased in the following order when compared
at the same ECe level: NaCl > CaCl2 > Na2SO4. Reduced enzyme activities in saline soils may
be due to the osmotic desiccation of microbial cells releasing intracellular enzymes which become vulnerable to attack by soil proteases, a
“salting-out” effect modifying the ionic conformation of the active site of the enzyme-protein, and specific ion toxicities causing nutritional
imbalances for microbial growth and subsequent enzyme synthesis.
The first plant organ to detect a limitation on the water supply is the root system. It has been shown that besides water and minerals, roots also send signals to
the leaves through the xylem sap, and the phytohormone abscisic acid is considered to be one of the major root-to-shoot stress signals. When the stress signal
reaches the leaves, it triggers stomatal closure and the plant shifts to a water-saving strategy. Hence, by adjusting stomatal opening, plants are able to control
water loss by reducing the transpiration flux, but they are concomitantly limiting the entrance of carbon dioxide (CO2). This will have direct and indirect effects
on the reduction of net photosynthesis and on the overall production of ROS by plants under drought stress. There are many studies that report on increased
ROS accumulation and oxidative stress under drought stress. In fact, under drought stress, ROS production is enhanced through multiple ways. For instance, the
limitation on CO2 fixation will reduce NADP+ regeneration through the Calvin cycle, hence provoking an over reduction of the photosynthetic electron transport
chain. In fact, during photosynthesis and under drought stress there is a higher leakage of electrons to O2 by the Mehler reaction. It was estimated that in
drought stressed wheat, the leakage of photosynthetic electrons to the Mehler reaction was increased by approximately 50% as compared to unstressed wheat.
An increase on thylakoid membrane electron leakage to O2 under drought stress was also seen in sunflower. However, it is quite difficult to assess the part of
ROS generated by the Mehler reaction to that generated by photorespiration. In fact, under drought stress the photorespiratory pathway is also enhanced,
especially when RuBP oxygenation is maximal due to limitation on CO2 fixation. The predominance of photorespiration on the oxidative load under drought
stress has been recently put forward. Noctor and collaborators have estimated that photorespiration is likely to account for over 70% of total H2O2 production
under drought stress conditions.
Abscisic acid (ABA) is a plant hormone involved in the response of plants to reduced water availability. Reduction of guard cell turgor by ABA diminishes the
aperture of the stomatal pore and thereby contributes to the ability of the plant to conserve water during periods of drought.
ROS are a group of free radicals, reactive molecules, and ions that are derived from O 2. It has been estimated that about 1% of O2 consumed by plants is diverted
to produce ROS
Further, limited CO2 availability due to closure of stomata during various environmental stresses such as salinity, drought favors the formation of 1O2. The life
time of 1O2 within the cell is probably 3 μs or less [30, 31]. A fraction of 1O2 has been shown to be able to diffuse over considerable distances of several hundred
nanometers (nm). 1O2 can last for 4 μs in water and 100 μs in a nonpolar environment [1]. 1O2 reacts with most of the biological molecules at near diffusion-
controlled rates [1]. It directly oxidizes protein, unsaturated fatty acids, and DNA [32]. It causes nucleic acid modification through selective reaction with
deoxyguanosine [33]. It is thought to be the most important species responsible for light-induced loss of photosystem II (PSII) activity which may trigger cell
death [34]. 1O2 can be quenched by β-carotene, α-tocopherol or can react with the D1 protein of photosystem II as target [29]. Due to spin restriction, molecular
O2 cannot accept four electrons at a time to produce H2O. It accepts one electron at a time and hence during reduction of O2 stable intermediates are formed in
the step-wise fashion [35]. O 2•− is the primary ROS formed in the cell which initiates a cascade of reactions to generate “secondary” ROS, either directly or
prevalently through enzyme- or metal-catalysed processes [36] depending on the cell type or cellular compartment. O 2•− is a moderately reactive, short-lived
ROS with a half-life of approx. 1 μs. O 2•− is a nucleophilic reactant with both oxidizing and reducing properties [37]. Anionic charge of O 2•− inhibits its
electrophilic activity toward electron-rich molecules. O 2•− has been shown to oxidize enzymes containing the [4Fe-4S] clusters (aconitase or dehydratase as
examples) [38] and reduce cytochrome C [39]. O 2•− can accept one electron and two protons to form H2O2.
Salinity-induced ROS disrupt normal metabolism through lipid peroxidation, denaturing proteins, and nucleic acids in several plant species