CN106165621B - γ-aminobutyric acid improves the purposes of the resistance to salt stress ability of corn - Google Patents
γ-aminobutyric acid improves the purposes of the resistance to salt stress ability of corn Download PDFInfo
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Abstract
Gamma aminobutyric acid improves the purposes of the resistance to salt stress ability of corn, is related to purposes of the gamma aminobutyric acid in corn growth.The invention aims to provide the new application of gamma aminobutyric acid.Maize under NaCl stress leaf oxidation is damaged gamma aminobutyric acid and enzymatic activity has regulating and controlling effect.Reduce the mda content in Maize Seedling Under Salt Stress blade.Reduce superoxide anion content in blade.Reduce superoxide dismutase content in blade.Contribute to the lifting of peroxidase activity.Improve the activity of the lower Antioxidant Enzyme Systems of NaCl stress.There is regulating and controlling effect to Maize Seedling Under Salt Stress nitrogen metabolism.Improve the soluble protein content of maize under NaCl stress blade and root system.Reduce maize under NaCl stress blade and acetone acid content, activity of glutamine synthetase, glutamate synthetase activity, glutamate dehydrogenase enzymatic activity and glutamate decarboxylase activity in root system.The present invention is used for corn and cultivates field.
Description
Technical field
The present invention relates to the purposes that γ-aminobutyric acid improves the resistance to salt stress ability of corn.
Background technology
Corn is the important grain in China and forage crop, is occupied an important position in national economy.Salinization soil
It is a kind of widely distributed low productive soil, is to cause one of main environment factor of crop failure in world wide.Salt stress meeting
The normal growth and development of crop is influenced, and corn is moderate salt density value type crop, salt stress can seriously affect the physiology of corn
Metabolism, so as to directly affect the growth and development of corn and yield and quality is formed.
γ-aminobutyric acid (GABA) is a kind of four carbon nonprotein amino acid being present in free state in various biologies,
It is highly soluble in water, and dependent on it, there are a variety of configurations in plant for this.Plant is such as joined to be contained in the seed of category, yam bean
GABA, also contains GABA in the rhizome of some plants in addition.GABA in animal body is almost existed only in nerve fiber,
It can participate in many metabolism processes, there is very high physiological activity, be important inhibitory neurotransmitter, and be also at present research compared with
For deep neurotransmitter.
GABA is a kind of chemical substance, it for the first time the artificial synthesized time be in 1883.Research shows that GABA exists
The normal intracerebral content of mammal is higher, but its physiological significance is still indefinite.Segal SA et al. confirm GABA to lactation
The nervous centralis of animal has universal inhibitory action.According to current research, the physiological activity of GABA is mainly manifested in following several respects：
1. calm nerve, antianxiety；2. reduce blood pressure；3. treat disease；4. reduce blood ammonia；5. improve brain vigor；6. promote ethanol generation
Thank.In addition GABA also plays an important role in healthcare.In the food industry, GABA can utilize prepared by food processing byproducts
GABA additives, using fermentation method biosynthesis GABA additives, development functionality dairy products, applied to bakery and
Applied in beverage etc..
The content of the invention
The invention aims to provide the new application of γ-aminobutyric acid.
γ-aminobutyric acid of the present invention improves the purposes of the resistance to salt stress ability of corn, and specially γ-aminobutyric acid is to salt stress
Lower maize leaf oxidative damage and enzymatic activity have regulating and controlling effect.
Further, Gabanergic reduces the mda content in Maize Seedling Under Salt Stress blade, so as to reduce
Damage to cell membrane.
Further, γ-aminobutyric acid processing can reduce superoxide anion content in Leaves of Maize Seedlings, external source γ-ammonia
Base butyric acid can reduce accumulation of the superoxide anion in blade.
Further, γ-aminobutyric acid reduces superoxide dismutase content in blade.
Further, γ-aminobutyric acid processing contributes to the lifting of peroxidase (POD) activity.GABA's is optimal dense
Spend for 0.5mmol/L.
Further, the Gabanergic of 0.5mmol/L improves the activity of the lower Antioxidant Enzyme Systems of NaCl stress, subtracts
The accumulation of few harmful substance, ensures the normal physiological metabolism of seedling.
γ-aminobutyric acid of the present invention improves the purposes of the resistance to salt stress ability of corn, and specially γ-aminobutyric acid is to salt stress
Lower corn seedling nitrogen metabolism has regulating and controlling effect.
Further, Gabanergic improves the soluble protein of the maize leaf and root system under NaCl stress growths
Content, alleviates influence of the NaCl stress to maize seedling roots.
Further, under NaCl stress, γ-aminobutyric acid processing can reduce acetone acid content in maize leaf and root system.
Further, γ-aminobutyric acid reduces the activity of glutamine synthetase of maize under NaCl stress blade and root system.
Further, γ-aminobutyric acid reduces the glutamate synthetase activity of maize under NaCl stress blade and root system.
Further, γ-aminobutyric acid reduces the glutamate dehydrogenase enzymatic activity of maize under NaCl stress blade and root system.
Further, γ-aminobutyric acid reduces the glutamate decarboxylase activity of maize under NaCl stress blade and root system.
Beneficial effects of the present invention：
The present invention provides the purposes that γ-aminobutyric acid improves the resistance to salt stress ability of corn.It has compared with prior art
There is following advantage：
1st, there is the activity of higher relative to other plant growth regulator, γ-aminobutyric acid；
2nd, γ-aminobutyric acid is the naturally occurring amino acid of organism, any toxicity is not present to human body, at corn
Residual hazard residual effect is not present after reason；
3rd, processing method is easy, workable.
Brief description of the drawings
Fig. 1 is influences of the 958 various concentrations GABA of Zheng Dan to mda content under salt stress；
Fig. 2 is influences of the east 253 various concentrations GABA of agriculture to mda content under salt stress；
Fig. 3 is influences of the 958 various concentrations GABA of Zheng Dan to superoxide anion content under salt stress；
Fig. 4 is influences of the east 253 various concentrations GABA of agriculture to superoxide anion content under salt stress；
Fig. 5 is 958 maize leaf superoxide anion Histochemical localizations of Zheng Dan；
Fig. 6 is 253 maize leaf superoxide anion Histochemical localization of eastern agriculture；
Fig. 7 is influences of the 958 various concentrations GABA of Zheng Dan to SOD activity under salt stress；
Fig. 8 is influences of the east 253 various concentrations GABA of agriculture to SOD activity under salt stress；
Fig. 9 is influences of the 958 various concentrations GABA of Zheng Dan to POD activity under salt stress；
Figure 10 is influences of the east 253 various concentrations GABA of agriculture to POD activity under salt stress；
Figure 11 is influences of the 958 various concentrations GABA of Zheng Dan to CAT activity under salt stress；
Figure 12 is influences of the east 253 various concentrations GABA of agriculture to CAT activity under salt stress；
Figure 13 is influences of the Zheng Dan 958GABA to blade soluble protein content under salt stress；
Figure 14 is influences of the east agriculture 253GABA to blade soluble protein content under salt stress；
Figure 15 is influences of the Zheng Dan 958GABA to root system soluble protein content under salt stress；
Figure 16 is influences of the east agriculture 253GABA to root system soluble protein content under salt stress；
Figure 17 is influences of the Zheng Dan 958GABA to blade pyruvic acid content under salt stress；
Figure 18 is influences of the east agriculture 253GABA to blade pyruvic acid content under salt stress；
Figure 19 is influences of the Zheng Dan 958GABA to root system acetone acid content under salt stress；
Figure 20 is influences of the east agriculture 253GABA to root system acetone acid content under salt stress；
Figure 21 is influences of the Zheng Dan 958GABA to blade activity of glutamine synthetase under salt stress；
Figure 22 is influences of the east agriculture 253GABA to blade activity of glutamine synthetase under salt stress；
Figure 23 is influences of the Zheng Dan 958GABA to root system activity of glutamine synthetase under salt stress；
Figure 24 is influences of the east agriculture 253GABA to root system activity of glutamine synthetase under salt stress；
Figure 25 is influences of the Zheng Dan 958GABA to blade glutamate synthetase activity under salt stress；
Figure 26 is influences of the east agriculture 253GABA to blade glutamate synthetase activity under salt stress；
Figure 27 is influences of the Zheng Dan 958GABA to root system glutamate synthetase activity under salt stress；
Figure 28 is influences of the east agriculture 253GABA to root system glutamate synthetase activity under salt stress；
Figure 29 is influences of the Zheng Dan 958GABA to blade glutamte dehydrogenase activity under salt stress；
Figure 30 is influences of the east agriculture 253GABA to blade glutamte dehydrogenase activity under salt stress；
Figure 31 is influences of the Zheng Dan 958GABA to root system glutamate dehydrogenase enzymatic activity under salt stress；
Figure 32 is influences of the east agriculture 253GABA to root system glutamate dehydrogenase enzymatic activity under salt stress；
Figure 33 is influences of the Zheng Dan 958GABA to blade glutamate decarboxylase activity under salt stress；
Figure 34 is influences of the east agriculture 253GABA to blade glutamate decarboxylase activity under salt stress；
Figure 35 is influences of the Zheng Dan 958GABA to root system glutamate decarboxylase activity under salt stress；
Figure 36 is influences of the east agriculture 253GABA to root system glutamate decarboxylase activity under salt stress.
Embodiment
Technical solution of the present invention is not limited to act embodiment set forth below, further includes between each embodiment
Any combination.
Embodiment one：Present embodiment γ-aminobutyric acid improves the purposes of the resistance to salt stress ability of corn, is specially
Maize under NaCl stress leaf oxidation is damaged γ-aminobutyric acid and enzymatic activity has regulating and controlling effect.
Embodiment two：The present embodiment is different from the first embodiment in that：γ-aminobutyric acid reduces the salt side of body
Compel the mda content in lower Leaves of Maize Seedlings, so as to reduce the damage to cell membrane.It is other with one phase of embodiment
Together.
Embodiment three：The present embodiment is different from the first embodiment in that：γ-aminobutyric acid reduces blade
Interior superoxide anion content, reduces accumulation of the superoxide anion in blade.It is other identical with embodiment one.
Embodiment four：The present embodiment is different from the first embodiment in that：γ-aminobutyric acid reduces blade
Interior superoxide dismutase content.It is other identical with embodiment one.
Embodiment five：The present embodiment is different from the first embodiment in that：γ-aminobutyric acid contributed to
The lifting of peroxidase activity, wherein the concentration of γ-aminobutyric acid is 0.5mmol/L during processing.Other and embodiment
One is identical.
Embodiment six：The present embodiment is different from the first embodiment in that：γ-aminobutyric acid improves NaCl
The activity of the lower Antioxidant Enzyme Systems of stress, reduces the accumulation of harmful substance.It is other identical with embodiment one.
Embodiment seven：Present embodiment γ-aminobutyric acid improves the purposes of the resistance to salt stress ability of corn, is specially
γ-aminobutyric acid has regulating and controlling effect to Maize Seedling Under Salt Stress nitrogen metabolism.
Embodiment eight：Present embodiment is unlike embodiment seven：γ-aminobutyric acid improves the salt side of body
Compel the soluble protein content of lower maize leaf and root system.It is other identical with embodiment seven.
Embodiment nine：Present embodiment is unlike embodiment seven：γ-aminobutyric acid reduces the salt side of body
Compel the acetone acid content in lower maize leaf and root system.It is other identical with embodiment seven.
Embodiment ten：Present embodiment is unlike embodiment seven：γ-aminobutyric acid reduces the salt side of body
Compel lower maize leaf and the activity of glutamine synthetase of root system, the glutamate synthetase activity of maize leaf and root system, corn
Blade and the glutamate dehydrogenase enzymatic activity and maize leaf of root system and the glutamate decarboxylase activity of root system.Other and specific reality
It is identical to apply mode seven.
To verify beneficial effects of the present invention, tests below is carried out：
1 materials and methods
1.1 test material
For trying corn variety：Salt-resistant type kind Zheng Dan 958, salt density value type kind east agriculture 253, select full seed, neat
Unanimously, the high and harmless seed of cleanliness, germination test is carried out before sowing.
Test plant growth regulator：γ-aminobutyric acid (GABA), purchased from sigma companies.
1.2 experimental design
Test and carried out in 2014,2015 in agricultural college of Northeast Agricultural University greenhouse.By select before water culture experiment sowing
Seed soak 24h at 25 DEG C of constant temperature, seed sowing is being placed with a diameter of 0.5m of vermiculite, depth is in the disk of 7cm,
Often 80 seeds of disk, afterwards in seed overlying vermiculite 2-3cm, optionally water daily.When seedling is grown to 21 heart of leaf, by seedling
Move to equipped with the 1/2 suddenly sink of Glan nutrient solution, air-breather 24h ventilates into sink, to ensure the aerobic of growth of seedling
Amount.One time of nutrition liquid, and periodic adjustment solution ph are replaced every 2d, it is growth of seedling optimum pH to ensure it, treats seedling
Handled and sampled when growing into 31 heart of leaf.3 repetitions of each test process.
1.2.1 Seed Germination Test designs
1.2.1.1NaCl the screening of concentration
Choose in the same size, the experiment seed of full seed, the surface of the seed with 0.1% HgCl2Solution disinfection 15min,
Rinsed with distilled water 3 times after taking-up, blotted afterwards with blotting paper repeatedly.Seed is immersed in 24h in clear water.Will be a diameter of
The culture dish of 150mm is cleaned, and is put into autoclave high temperature and is sterilized 30 minutes.2 layers of filter paper are put into culture dish, and respectively
Add (1) clear water (CK), (2) 50mmol/L NaCl (AT1), (3) 100mmol/L NaCl (AT2), (4) 150mmol/L
NaCl (AT3), (5) 200mmol/L NaCl (AT4) solution 20ml.Seed embryo after immersion is upward and equidirectional be placed in training
Support in ware, 80 are put in each culture dish, 1 layer of filter paper is put on seed, water conservation moisturizing, repeats three times.Growth cabinet temperature is
28/22 DEG C, daily lighting delay number is 12h, and number is 12h when dark, relative humidity 60%, intensity of illumination 4000lux.Main root
It is germination standard to grow when reaching seed length half, counts germinative number daily.
1.2.1.2 various concentrations GABA regulates and controls Seed Germination Test
Choose in the same size, the experiment seed of full seed, the surface of the seed with 0.1% HgCl2Solution disinfection 15min,
Rinsed with distilled water 3 times after taking-up, blotted afterwards with blotting paper repeatedly.Seed is immersed in 24h in clear water.Will be a diameter of
The culture dish of 150mm is cleaned, and is put into autoclave high temperature and is sterilized 30 minutes.2 layers of filter paper are put into culture dish, and respectively
Add (1) clear water (CK), (2) 100mmol/L NaCl (AN), (3) 100mmol/L NaCl+0.25mmol/L GABA
(ANG1), (4) 100mmol/L NaCl+0.5mmol/L GABA (ANG2), (5) 100mmol/L NaCl+1mmol/L GABA
(ANG3), (6) 100mmol/L NaCl+2mmol/L GABA (ANG4).Seed embryo after immersion is upward and equidirectional put
In culture dish, 80 are put in each culture dish, 1 layer of filter paper is put on seed, water conservation moisturizing, repeats three times.Artificial climate box temperature
Spend for 28/22 DEG C, daily lighting delay number is 12h, and number is 12h when dark, relative humidity 60%, intensity of illumination 4000lux.
It is germination standard that main root, which is grown when reaching seed length half, counts germinative number daily.
1.2.2 growth of seedling water culture experiment designs
1.2.2.1NaCl the screening of concentration
When growth of maize is to 31 heart of leaf, handled.Before processing replaces once Huo Gelan nutrient solutions, adds afterwards
NaCl so that the concentration of NaCl in the solution respectively reaches 0mmol/L (CK), 50mmol/L (BN1), 100mmol/L (BN2),
150mmol/L (BN3), 200mmol/L (BN4) and 250mmol/L (BN5).48h is measured by sampling after adding sodium chloride.Blade takes
2nd leaf, it is stand-by with tinfoil ladle sample, -80 DEG C of preservations.
1.2.2.2GABA the screening test of concentration
When growth of maize is to 31 heart of leaf, handled.Before processing replaces once Huo Gelan nutrient solutions, adds afterwards
GABA so that the concentration of GABA in the solution respectively reaches 0mmol/L (CK), 0.25mmol/L (BNG1), 0.5mmol/L
(BNG2), 1.0mmol/L (BNG3), 2.0mmol/L (BNG4).After adding the 24h of GABA, NaCl is added in solution in three times,
So that ultimate density is 150mmol/L, as 0h when NaCl concentration is 150mmol/L, add 0 after NaCl, 12,24,36,
48h, is measured by sampling.Blade takes the 2nd leaf, root system take it is well-grown, it is stand-by with tinfoil ladle sample, -80 DEG C of preservations.
1.2.2.3GABA regulate and control growth of seedling experiment
When growth of maize is to 31 heart of leaf, handled.Before processing replaces once Huo Gelan nutrient solutions, adds afterwards
GABA so that the concentration of GABA in the solution reaches 0 and 0.5mmol/L.After adding the 24h of GABA, added in three times in solution
NaCl so that ultimate density is 0 and 150mmol/L, as 0h when NaCl concentration is 150mmol/L, and four processing are respectively
0mmol/L GABA+0mmol/L NaCl (CK), 0.5mmol/L GABA+0mmol/L NaCl (G), 0mmol/L GABA+
150mmol/L NaCl (N), 0.5mmol/L GABA+150mmol/L NaCl (NG).Add NaCl after 0,12,24,36,48h,
It is measured by sampling.Blade takes the 2nd leaf, root system take it is well-grown, it is stand-by with tinfoil ladle sample, -80 DEG C of preservations
1.3 testing indexs and method
1.3.1 oxidative damage and activities of antioxidant enzymes index
Malonaldehyde 1.3.1.1 (MDA) content
Thiobarbituricacidα- method measures.
1.3.1.2 superoxide anion (02 -) content
Measured using oxyammonia oxidizing process.
1.3.1.3 superoxide dismutase (SOD) is active
Measured with NBT reduction methods.
1.3.1.4 peroxidase (POD) is active
Using guaiacol method.
1.3.1.5 catalase (CAT) is active
5.73ml PBS solution+0.2ml enzyme liquids, measure pipe boiling water bath 10min, control tube not water-bath.Will after 10min water-baths
After 25 DEG C of water-bath 5min of control and measure pipe, 0.07ml H are added by pipe2O2Reaction solution, often adds i.e. timing after a pipe, and rapidly
Pour into quartz cuvette, colorimetric under 240nm, every 1min readings 1 time, reading 3 times altogether.
1.3.2 nitrogen metabolism index of correlation measures
1.3.2.1 soluble protein content
Coomassie Brilliant Blue measures.
1.3.2.2 glutamte dehydrogenase (GAD) is active
Using GABA quantitative colorimetrs measure glutamate decarboxylase (GAD) activity, 1 μm of ol GABA is generated with every 30min and is made
For an enzyme activity unit.
1.3.2.3 glutamine synthelase (GS) is active
With reference to the method for Oaks etc., make standard curve with γ-glutamyl hydroxamic acid (GHA), measure GS vigor, enzyme activity
Power is with a μm ol NADHg-1FM·min-1Represent.
1.3.2.4 glutamate synthetase (GOGAT) is active
The enzyme liquid of glutamate synthetase (GOGAT) extracts same GS, and determination of activity is with reference to Singh and Srivastava[166]'s
Method, enzyme activity μm ol NADHg-1FM·min-1Represent.
1.3.2.5 glutamte dehydrogenase (GDH) is active
The enzyme liquid of glutamte dehydrogenase (GDH) extracts same GS, and determination of activity is with reference to the method for Lin and Kao, enzyme activity μ
mol NADH·g-1FM·min-1Represent.
1.3.2.6 acetone acid content
0.5g fresh plant tissues are weighed in mortar plus appropriate 8% trichloroacetic acid, being carefully ground into homogenate, then with 8% 3
Monoxone washes 25ml volumetric flasks, is settled to scale.Bottle stopper is stoppered, shaking extraction, stands 30min.Take about 10ml homogenates from
The heart (4000r/min) 10min, supernatant are spare.1.0ml supernatants are taken in scale test tube, adds 8% trichloroacetic acids of 2ml, adds
1.0ml 0.1%2,4- dinitrophenylhydrazine liquid, shake up, then add 5.0ml 1.5mol/L, and NaOH solution shakes up colour developing, after 10min
The colorimetric under 520nm wavelength, records absorbance, and the acetone acid content of measure pipe is checked on standard curve.
2 results and analysis
2.1 external source GABA coerce NaCl the regulation and control of lower maize leaf oxidative damage and enzymatic activity
2.1.1 to the influence of mda content
Malonaldehyde is the product of Lipid peroxidation metabolism, has detrimental effect to cell membrane.As illustrated in fig. 1 and 2 (in figure ◆ represent
CK, represent BN, ▲ represent BNG1, × represent that BNG2, ■ represent BNG3, ● represent BNG4), 958 control treatments of Zheng Dan the third two
Aldehyde (MDA) does not change significantly in 0-48h, the situation of mda content rising all occur in other processing.Wherein BN
Handle mda content to handle higher than other, this malonaldehyde that shows GABA and can reduce in Maize Seedling Under Salt Stress blade contains
Amount, so as to reduce the damage to cell membrane.When coercing 48h, BNG2 treatment effects are better than other GABA processing, and and its
His GABA processing differences are notable.In 0-48h, there is larger fluctuation in eastern 253 leaf malondialdehyde content of agriculture.The third the two of control
Aldehyde change is smaller, and BN processing content in 0-24h rises, and declines afterwards, and 36-48h obvious content occurs and rises again.
All processing of GABA generally show content in 0-12h and rise, and mda content declines afterwards, in 24-48h, MDA
Content persistently rises.In 48h, the mda content of BNG2 processing is less than others GABA processing, and significant difference.As a result table
Bright, due to breediness, mda content amplitude of variation is smaller in body when being subject to NaCl to coerce by Zheng Dan 958, and eastern agriculture 253 changes
Acutely.After GABA processing, the mda content of two kinds declines with respect to BN processing, this shows that GABA can reduce stress lower film
The generation of lipoperoxide, reduces the infringement to cell.In the GABA concentration for examination, BNG2 (0.5mmol/L) treatment effect
It is better than other processing, this shows that the Auto-regulator of suitable concentration has significant impact for regulating effect.
2.1.2 to the influence of superoxide anion content
First generation is exactly superoxide anion when oxygen produces oxygen radical in plant, itself is had malicious
Property, other oxygen radicals can be produced after carrying out a series of reaction, to the toxic effect of cell.Understood by Fig. 3 and 4 (in figure
◆ represent that CK, represent BN, ▲ represent BNG1, × represent that BNG2, ■ represent BNG3, ● represent BNG4), with stress time
Increase, Zheng Dan 958 compares blade superoxide anion content and keeps stablizing, and ascendant trend then occur in other processing.BN processing exists
Handled in 24-48h higher than other, and it is notable with other processing differences during 48h.GABA processing superoxide anion contents are less than at BN
Reason, wherein BNG2 processing blade superoxide anion content are less than other GABA processing, and in 36h and 48h and other GABA processing
Significant difference.The control treatment of eastern agriculture 253 in stress time superoxide anion content relatively stablize, less than it is all processing and
Each sample point difference is notable.BN processing slightly declines in 0-12h superoxide anions content, and what rear appearance significantly rose shows
As, and it is higher than all processing, and significant difference in 36h and 48h superoxide anions content.GABA respectively handles superoxide anion content
" W " type is presented in variation tendency, and highest numerical value is presented in 24h.Wherein BNG2 processing content is less than other GABA processing, still
Difference is not notable.The result shows that when not coerced, superoxide anion content tends towards stability in Leaves of Maize Seedlings, works as salt stress
When then there is fluctuation by a relatively large margin.GABA processing can reduce superoxide anion content in blade, and BNG2 treatment effects are best.
Between two kinds, the BN processing superoxide anion changes of contents of Zheng Dan 958 is relatively steady, and the amplitude that eastern agriculture 253 changes is larger,
Content during 48h does not occur steady or downward trend.In GABA processing, go out after the superoxide anion content 24h of eastern agriculture 253
Now obvious downward trend, Zheng Dan 958 is then slightly decline or steady, so GABA is for eastern 253 blade superoxide anion of agriculture
The mitigation of accumulation is better than Zheng Dan 958.
2.1.3 maize leaf superoxide anion Histochemical localization
Superoxide anion Histochemical localization technology is by O2 ·-Distribution and content visualization in plant body.It is blue in figure
Pointing object represents content and distribution of the superoxide anion in blade.In two corn varieties, control treatment pointing object is less, BN
Processing pointing object is most and is distributed in whole blade.External source GABA can reduce accumulation of the superoxide anion in blade, BNG2
Treatment effect is best.The BNG2 processing pointing object of two kinds is minimum, and is distributed in the middle and back of blade more.At other GABA
Reason pointing object compared with BN processing has been reduced, but effect unobvious, and pointing object is distributed in whole blade (Figures 5 and 6).
2.1.4 to the influence of superoxide dismutase activity
SOD can be disproportionated two O2 ·-Free radical becomes H2O2And O2, this process for protection cell it is most important.Such as Fig. 7
With shown in 8 (in figure ◆ represent CK, represent BN, ▲ represent BNG1, × represent BNG2, ■ represent BNG3, ● represent BNG4), with
The increase of stress time, except for the control, larger ripple occurs in superoxide dismutase (SOD) activity of two corn varieties
It is dynamic.The 0-12h of NaCl stress, due to the self-defense characteristic of plant, there is obvious rise, BNG2 in two kind SOD activity
For SOD activity with BNG4 processing higher than other processing, difference is not notable each other, but notable with other processing differences.After stress
12-24h, as active oxygen radical in the reduction of plant in-vivo content or tends towards stability, the SOD activity of Zheng Dan 958 is in now
Drop trend, eastern 253 part of agriculture processing SOD activities presents go out stable state, but BNG2 and BNG1 processing then shows active rising
Trend.After the 24-48h of stress, due to the persistent accumulation of active oxygen, the SOD sustained activities of Zheng Dan 958 raise, in order to clear
Except activity in vivo oxygen, wherein BNG2 treatment effects are best.In addition to BNG2 processing, other are handled after 36-48h is coerced for eastern agriculture 253,
SOD activity declines.Wherein BN is active minimum processing, and except for the control notable with other processing differences, BNG2 activity highests,
It is and notable with other processing differences.The result shows that two corn varieties are different for NaCl stress responses sensitivity, Zheng Dan
958 quickly can respond stress, and eastern agriculture 253 then responds slowly, this may be related to stress resistance difference with the two.
2.1.5 to the influence of peroxidase activity
H2O2Toxic action can be produced to cell, POD is considered as H main in SCAVENGING SYSTEM OF ACTIVATED OXYGEN2O2Scavenger enzyme.
Knowable to Fig. 9 and 10 (in figure ◆ represent that CK, represent BN, ▲ represent BNG1, × represent BNG2, ■ expression BNG3, ● represent
BNG4), peroxidase (POD) activity of two corn variety controls changes unobvious with the increase of stress time, other
There is obvious change in processing.The POD activity of two kinds, which is presented, first rises the trend for declining afterwards, rising again afterwards.
NaCl coerces initial stage, due to the defense mechanism of corn itself, internal POD activity can be raised, in order to remove H2O2, at GABA
Reason contributes to the lifting of POD activity, and wherein BNG2 treatment effects are the most obvious.The 12h after stress, what two kind BNG2 were handled
POD activity is handled higher than other, and significant difference.With stabilization of the harmful substance in corn seedling body, the 12- after stress
Downward trend is presented in 36h, POD activity, the active highest of BNG2 processing in all processing of this process.36h after stress
For the part processing POD activity of eastern agriculture 253 less than control, this is probably because NaCl stress causes one to the physiological mechanism of seedling
Fixed damage, makes the processes such as the synthesis of enzyme be obstructed.The 48h after stress, with the accumulation of the materials such as active oxygen, in corn seedling body
POD activity increase, the BN of two of which kind processing activity is minimum, the active highest of BNG2 processing, but with other GABA
Processing difference it is not notable.The result shows that in the corn seedling of stress is not affected by, POD is activity stabilized, beautiful when being forced
Meter You Miao can start host defence mechanism, resistance damage.GABA contributes to the lifting of POD activity, the GABA of wherein BNG2 processing
Concentration (0.5mmol/L) effect is ideal.
2.1.6 to the influence of catalase activity
CAT can be by H2O2It is decomposed into H2O and O2, so as to reduce H2O2Content.With the increase of stress time, except control
Outside, catalases (CAT) activity of other processing is presented first raise after downward trend, the 24h activity highests after stress.Zheng
In single 958, the CAT activity of BN processing is less than the processing containing GABA, and wherein BNG2 is better than other GABA processing, and in 12 Hes
24h and others GABA processing differences are notable.After coercing 48h, BNG2 processing activity is higher than other processing, but difference is not notable.
In eastern agriculture 253, BNG2 processing still effect is better than other processing, and 36h after stress and 48h are significantly higher than its elsewhere
Reason, see Figure 11 and 12 (in figure ◆ represent CK, represent BN, ▲ represent BNG1, × represent BNG2, ■ represent BNG3, ● represent
BNG4).For the CAT activity of the 48h after stress, BN processing less than the activity of control, this is probably because stress causes certain of corn
A little physiological barriers, cause the route of synthesis of enzyme to be obstructed.
In summary data are understood, the GABA of 0.5mmol/L can improve the activity of the lower Antioxidant Enzyme Systems of NaCl stress, subtract
The accumulation of few harmful substance, ensures the normal physiological metabolism of seedling.Therefore, it is optimal for establishing of seedling morphology also to demonstrate this concentration
The conclusion of concentration, afterwards experiment are carried out using 0.5mmol/L GABA.
2.2 external source GABA coerce NaCl the regulation and control of lower corn seedling nitrogen metabolism
2.2.1 to the influence of soluble protein content
2.2.1.1 to the influence of leaf soluble protein content
Soluble protein is important osmotic adjustment and nutriment, their increase and accumulation can improve cell
Water holding capacity, plays a protective role the living matter and biomembrane of cell.Knowable to Figure 13 and 14 (represents CK,Represent
G,Represent N,Represent NG), with the increase of stress time, the leaf soluble protein content content of two corn varieties is presented
First raise and decline afterwards, afterwards elevated trend again.In Zheng Dan 958, the order of soluble protein content is G ＞ in four processing
CK ＞ NG ＞ N.This shows that GABA can significantly improve the soluble protein being not affected by 958 blades of Zheng Dan of NaCl stress and contain
Amount, increasing degree 12.61%-38.89%, in addition to the 24h after stress, other times are notable with contrast difference.With compareing
Compare, NaCl can significantly reduce the content of the soluble protein in 958 blades of Zheng Dan, and the amplitude of reduction is 28.91%-
37.39%, all sample time differences are notable.Under NaCl stress, soluble egg in blade can be significantly improved using GABA
Bai Hanliang, alleviates the phenomenon that soluble protein content declines caused by NaCl is coerced.Soluble protein contains in NG processing blades
Amount is higher than N processing, and increased amplitude is 17.42%-30.77%, and in addition to the 24h after stress, other sample time differences are equal
Significantly.For soluble protein content higher than compareing, the ratio of raising is 17.46%-34.37% in the G processing blades of eastern agriculture 253,
In addition to the 48h of stress, other times significant difference.NaCl stress significantly reduces soluble protein content in blade, with
The administration of GABA, the protein content of NG processing is significantly higher than N processing, 14.96%-78.18% is added, after stress
24h differences are not significantly outer, other sample time significant differences.Two kinds are compared, 253 corn seedling pair of eastern agriculture after stress
The response of GABA is more strong, and soluble protein content excursion is more than Zheng Dan 958 in its blade.
2.2.1.2 the influence to root system soluble protein
With the increase of stress time, the soluble protein contents of two kind root systems, which is showed, identical with blade to become
Gesture.The root system soluble protein content of two kinds is ordered as G ＞ CK ＞ NG ＞ N, this shows that GABA can improve normal growth bar
The soluble protein content of part and the root system under NaCl stress growths, alleviates influence of the NaCl stress to maize seedling roots.
In Zheng Dan 958, G is handled with the soluble protein content compareed in addition to 12h differences are not notable, other times significant difference.Often
The NG processing soluble protein contents of a sample time are above N processing, and increased scope is being coerced between 12%-45.16%
12h, 24h and 36h significant difference after compeling.The G processing solubles protein content of eastern agriculture 253 is higher than control treatment, but each
Sample time difference is not notable, and the amplitude of raising is between 4.25%-12.09%.The soluble protein content of NG processing and N processing
In addition to 0h differences are not notable, other sample times equal significant difference, and NG processing handles the amplitude improved in 18.84%- compared with N
Between 48.75%, see Figure 15 and 16 (represent CK,Represent G,Represent N,Represent NG).
2.2.2 to the influence of acetone acid content
2.2.2.1 to the influence of blade acetone acid content
Pyruvic acid participates in one of intermediate product of whole organism analytic metabolism.Pyruvic acid can pass through acetyl-CoA and tricarboxylic
The mutual inversion of phases between sugar, fat and amino acid in vivo is realized in acid circulation, and therefore, pyruvic acid is contacted in the metabolism of three major nutrient
In play important pivotal role.As shown in FIG. 17 and 18 (represents CK,Represent G,Represent N,Represent NG), with the side of body
Compel the increase of time, ascendant trend is substantially presented in two corn variety blade acetone acid contents.In Zheng Dan 958, G processing blades
Interior acetone acid content is less than control, and in addition to both 0h differences are not notable, other times difference is notable.Compared with the control, NaCl
Stress can cause acetone acid content in blade to rise, and the content of pyruvic acid in blade can be reduced after GABA processing, after stress
It is lower by 23.85% than N processing that 36h, NG handle acetone acid content.In addition to the NG processing of stress 0h and N processing differences be not notable, other
Sample time, two processing differences were notable.The G processing acetone acid content of eastern agriculture 253 is below compareing in 12,24 and 36h, and
Significant difference.The acetone acid content of all sample point NG processing is below N processing, and the amplitude of reduction is 7.14%-37.5%, is removed
0h differences are not notable, other times significant difference.
2.2.2.2 to the influence of root system acetone acid content
Two kind root system acetone acid contents increase with the increase of stress time, the 36h content highests after stress, it
After downward trend is presented, see Figure 19 and 20 (represent CK,Represent G,Represent N,Represent NG).In Zheng Dan 958, at G
In 0h, higher than control, the content of other sample times is below compareing acetone acid content reason, and after stress 12h and 24h with
Contrast difference is notable.N processing acetone acid contents after stress after 12,24,36 and 48h are shown higher than control with contrast difference
Write.In 12-48h after stress, NG processing acetone acid content is less than N processing, and the amplitude of reduction is 8.25%-29.33%.In east
In agriculture 253, the G processing acetone acid content of the 12-48h after stress is less than control treatment, and significant difference；NG processing pyruvic acid contains
Amount is less than N processing, and the 12-48h significant differences after stress.The result shows that GABA processing under normal growing conditions and
Acetone acid content in maize root system can be reduced under NaCl stress, but under NaCl stress, GABA reduces the effect of acetone acid content
Fruit is better than normal growing conditions.Two kinds compare, and GABA is more notable to eastern 253 regulating effect of agriculture.
2.2.3 to the influence of glutamine synthelase
2.2.3.1 the influence to blade glutamine synthelase
Glutamine synthelase (GS) is the key enzyme of plant nitrogen metabolism, it is catalyzed glutamic acid in glutamic acid synthesis circulation
With NH3Glutamine is formed, participates in the metabolism of plant nitrogenous compound.As shown in figs. 21 and 22 (represents CK,Represent
G,Represent N,Represent NG), glutamine synthelase (GS) activity is raised with the increase of stress time.In Zheng Dan 958
In, the GS activity of N processing respectively than compare it is high by 0.5%, 23.67%, 56.81%, 43.23% and 42.72%, and except coercing
0h differences are not notable after compeling, other times significant difference.The GS activity of NG processing is less than N processing, and 12,24,36 after stress
With 48h respectively it is higher by 11.96% than N processing, 42.43%, 17.28% and 7.42%, and significant difference.In eastern agriculture 253, at G
The GS activity of 24, the 36 and 48h after stress is managed less than compareing, it is lower by 38.43%, 24.84% and 10.77% than compareing respectively,
It is and significantly not outer with contrast difference.NaCl stress can cause the GS activity rises in eastern 253 blade of agriculture, can be reduced using after GABA
GS activity.GS activity after 0,12,24,36 and 48h of the NG processing after stress is below N processing, distinguish low 9.85%,
16.07%th, 39.74%, 43.27% and 49.58%, in addition to 0h differences are not notable, other times significant difference.The result shows that G
Processing can reduce blade GS activity, and the lower GABA that adds of NaCl stress can reduce the GS activity of blade.Compared with N processing, Zheng
The reduction ratio of single 958 NG processing reduces afterwards as stress time first increases, and eastern agriculture 253 is then increased over time and risen
Height, the effect that the eastern agriculture 253 of GABA regulation and control reduces blade GS activity are better than regulation and control Zheng Dan 958, and regulation and control of the GABA under stress
Enzymatic activity effect is better than under normal growing conditions.
2.2.3.2 the influence to root system glutamine synthelase
With the increase of stress time, the GS activity increases in 253 root system of Zheng Dan 958 and Dong Nong.In Zheng Dan 958,
NaCl stress after N processing GS activity higher than control, each sample point respectively than compare it is high by 3.59%, 31.31%,
30.52%th, 35.09% and 14.8%, in addition to 0h differences are not notable, other significant differences.The GS activity of NG processing is less than at N
Reason, and 12,24,36 after stress and 48h significant differences.In eastern agriculture 253, in addition to 0 after stress and 48h, other samplings
The GS activity of the G processing of point is less than control treatment.The GS activity of 0h of the N processing after except stress is high less than control, other times
In control, and significant difference.NG processing is below N processing in the GS activity of all sample points, distinguish low 15.61%,
5.47%th, 18.35%, 32.53% and 8.00%, and 24,36 and 48h significant differences (Figure 23 and 24, tables in figure after stress
Show CK,Represent G,Represent N,Represent NG).
2.2.4 to the influence of glutamate synthetase
2.2.4.1 the influence to blade glutamate synthetase
Glutamate synthetase (GOGAT) is important enzyme during nitrogen metabolism, it is main in the synthesis circulation of glutamic acid
It is catalysis glutamine and α-ketoglutaric acid generation glutamic acid.As shown in figs. 25 and 26 (represents CK,Represent G,Represent
N,Represent NG), glutamate synthetase (GOGAT) activity of two kind blades rises with the increase of stress time.
In Zheng Dan 958, the GOGAT activity of N processing is all higher than control, and significant difference in all sample times.The GOGAT of G processing lives
Property after stress 0 and 48h higher than control, other times less than control.The GOGAT activity of NG processing is below in all sample points
N processing, and it is notable with N processing differences.In eastern agriculture 253,12-36h of the GOGAT activity after stress of G processing less than control,
The GOGAT activity of N processing is in 12-48h higher than control.The enzymatic activity of NG processing is less than N processing, and the 0h after except stress
Outside, other times are notable with contrast difference.The result shows that GABA is for GOGAT activity in blade under conditions of normal growth
The effect of reduction is smaller, and rise effect of the NaCl stress for GOGAT activity is notable, can be bright using GABA under NaCl stress
Aobvious reduction GOGAT activity, lower response of the corn seedling to GABA of this explanation NaCl stress are more effective.
2.2.4.2 the influence to root system glutamate synthetase
Downward trend is presented in 0-12h of the GOGAT activity of Zheng Dan 958 after stress, and ascendant trend is presented afterwards.After stress
0h, four no obvious differences of processing.The GOGAT activity of 12-48h after stress, G processing is being coerced less than control
12,24 afterwards and 48h significant differences.The GOGAT activity of N processing is higher than control, and in addition to 0h after stress, other times are with compareing
Significant difference.The GOGAT activity of NG processing be less than N processing, respectively than N handle it is low 3.43%, 6.90%, 21.22%,
14.66% and 18.14%, and it is notable in 24-48h and the N processing difference of stress.In eastern agriculture 253, with the increasing of stress time
Add, the GOGAT activity rises in root system.The GOGAT activity of NG processing is less than N processing, and the 12-48h after stress and N processing is poor
It is different significantly (Figure 27 and 28, represents CK in figure,Represent G,Represent N,Represent NG).
2.2.5 to the influence of glutamte dehydrogenase
2.2.5.1 the influence to blade glutamte dehydrogenase
Two corn varieties glutamte dehydrogenase (GDH) activity is improved with the increase of stress time, see Figure 29 and
30 (represents CK,Represent G,Represent N,Represent NG).In Zheng Dan 958, compared with control treatment, G processing reduces
The activity of GDH, compared with the control, 12-48h of the N processing after stress significantly improve the activity of GDH, are carried respectively than control
It is high by 27.79%, 25.80%, 57.12% and 149.59%, and significant difference compared with the control.Handled using the NG of GABA
GDH activity be less than N processing, the 0-48h after NaCl is coerced respectively than N handle it is low 3.17%, 16.93%, 8.49%,
28.31% and 32.17%, in addition to 0h and N processing differences be not notable, other sample times equal significant difference.In eastern agriculture 253,
0h after stress, four processing GDH activity are without obvious difference.The GDH activity of G processing is less than control treatment, but is only coercing
Two processing differences of 12h are notable after compeling, other times GDH activity change unobvious.Compared with the control, N processing 12- after stress
The GDH activity of 48h is higher than control, and 12-48h and contrast difference are notable after stress.GABA processing under stress is obvious
GDH activity is reduced, it is lower by 12.97%, 11.52%, 32.31% and 38.94% than N processing respectively in 12-48h, and difference is shown
Write.
2.2.5.2 the influence to root system glutamte dehydrogenase
Glutamte dehydrogenase (GDH) activity in 958 root systems of Zheng Dan is improved with the increase of stress time.Coerced in NaCl
The GDH activity of 12-36h after compeling, G processing is less than control, and notable in 12,36 and 48h and contrast difference.NaCl stress carries
GDH activity in high 958 root systems of Zheng Dan, 12-48h after stress respectively than compare it is high by 27.60%, 12.99%,
44.04% and 97.81%.The activity for applying the NG processing of GABA compares display downward trend compared with N processing, is coerced compared with N processing
The GDH activity of 12-48h after compeling reduces by 20.64%, 10.20%, 31.11% and 23.84%, and significant difference respectively.
In eastern agriculture 253, in addition to 0h after stress, the GDH activity of other times G processing is less than control, and significant difference.After stress
In 12-48h, N handles GDH activity apparently higher than control, and significant difference.The GDH activity of NG processing is less than N processing, and is coercing
After compeling 12-48h and N processing differences significantly (Figure 31 and 32, represents CK in figure,Represent G,Represent N,Represent NG).
2.2.6 to the influence of glutamate decarboxylase
2.2.6.1 the influence to blade glutamate decarboxylase
Glutamate decarboxylase (GAD) can be catalyzed glutamic acid decarboxylase generation γ-aminobutyric acid (GABA).As seen in figs. 33 and 34
(represents CK,Represent G,Represent N,Represent NG), in Zheng Dan 958, the work of four processing glutamate decarboxylases (GAD)
0h does not have obvious difference after stress between property.Compared with the control, G processing has declined GAD activity.The GAD of N processing lives
Property 12-48h after stress significantly higher than control, respectively than compareing high by 42.98%, 72.61%, 164.54% and
188.54%, and significant difference.The NG processing GAD activity that application of GABA is less than N processing, the 12-48h after stress, difference
It is lower by 20.69%, 21.28%, 45.16% and 25.68% than N processing, and significant difference.In eastern agriculture 253, the GAD of G processing lives
Property 12-48h after stress less than control, and in 12,36 and 48h significant differences.12- of the GAD activity of N processing after stress
48h is higher than control, and significant difference.With the administration of the lower GABA of stress, the GAD activity that NG is handled compared with N processing is shown
The reduction of work, the 12-48h after stress are respectively lower than N processing 19.79%, 12.04%, 45.45% and 7.00%.The result shows that
Under normal growing conditions, raising influences of the GABA for GAD activity is smaller, but can significantly be dropped using GABA under NaCl stress
GAD activity in low two kind blades.
2.2.6.2 the influence to root system glutamate decarboxylase
The trend risen is presented with the time of stress in glutamate decarboxylase (GAD) activity of two corn variety root systems,
36h activity highests after stress.In Zheng Dan 958, without obvious poor between the GAD activity of tetra- processing of 0h after stress
It is different.Activity of the G processing in 12-48h is less than control.N processing activity 12-48h after stress significantly higher than control, is compared respectively
According to it is high by 36.75%, 64.68%, 114.94% and 124.62%.The GAD activity for including the NG processing of GABA is significantly lower than N
Handle, the 12-48h after stress is lower by 15.23%, 18.93%, 38.40% and 29.41% than N processing respectively, and poor with N processing
It is different notable.In eastern agriculture 253, the GAD activity of G processing is less than control, and the 12-48h significant differences after stress.NaCl can be carried
GAD activity in high root system, the GAD activity of the 12-48h after stress, N processing is apparently higher than control, and significant difference.Stress
The administration of lower GABA can reduce GAD activity, the 12-48h after stress, NG processing respectively lower than the GAD activity of N processing 17.73%,
29.05%th, 36.63% and 20.88%, and significant difference (Figure 35 and 36, represents CK in figure,Represent G,Represent N,Represent NG).
Claims (1)
- A kind of 1. method for improving the resistance to salt stress ability of corn, it is characterised in that：The corn variety selects salt density value type kind Eastern agriculture 253, carries out it in γ-aminobutyric acid (GABA) regulation and control seed and sprouts processing and γ-aminobutyric acid (GABA) regulation and control seedling Growth process,γ-aminobutyric acid (GABA) the regulation and control seed sprouting processing includes：Choose in the same size, the experiment kind of full seed Son, the surface of the seed with 0.1% HgCl2Solution disinfection 15min, is rinsed 3 times, uses blotting paper afterwards repeatedly after taking-up with distilled water Blot；Seed is immersed in 24h in clear water again, the culture dish of a diameter of 150mm is cleaned and is put into the sterilization of autoclave high temperature 30 minutes；2 layers of filter paper are put into culture dish, add the GABA immersions of the NaCl and 0.5mmol/L of 100mmol/L；Will immersion Seed embryo afterwards it is upward and it is equidirectional be placed in culture dish, 80 are put in each culture dish, 1 layer of filter paper, water conservation are put on seed Moisturizing, repeats three times；Growth cabinet temperature is 28/22 DEG C, and daily lighting delay number is 12h, and number is 12h when dark, relatively wet Spend for 60%, intensity of illumination 4000lux；γ-aminobutyric acid (GABA) the regulation and control growth of seedling processing includes：When growth of maize is to 31 heart of leaf, located Reason；Before processing replaces once Huo Gelan nutrient solutions, adds GABA afterwards, the concentration of GABA in the solution is reached 0.5mmol/L, After adding the 24h of GABA, NaCl is added in solution in three times, it is 150mmol/L to make its ultimate density.
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