Title: ????? ???? Salinity stress
1 ????? ???? Salinity stress
- ????? ?? ???? ?????
- ???? ??? ?? ????? ??? ??????
- ?? ????? ??? ???? ???? ????
2????? ????
- ????? ????? ???? ????? ??? ?? ???? ????? ?????
??? ?? ??? ??? ? ?? ????? ???? ???? ?? ??? ?? ??
??? ???? ? ????? ???? ??? ? ?? ???? ?? ??????
???? ???. - - ????? ???? ?? ?? ?? ???? ?? ???? ????? ??? ???
??????? ??? ?? ???? ???? ??? ???? ?? ClNa ??? - - ?????????? ??? ?? ????? ClNa ???.
3????? ??? ??? ?????
- CaCl2 ? MgCl2 ? MgSo4? Na2So4 ? ClNa ? CaSo4?
Ca(HCo3)2? NaHCo3 - ?? ???????
- CaCl2 MgCl2
NaCl - Ca CaSo4 Mg Na
NaHco3 - Ca(HCo3)2 MgSo4
Na2co3 - ??? 6 ?? ?? ?? ??? ???? ??? ???? ????? ???????
????? ??????? ????? ????? - ?? ????? 12 ?? ?? ????? ???? ??? ?? ???? ?? ?????
?? 50 ?? ?? ????? ?? ???? ???. ??? ??? ????? ??
???? ??????? ???? ?? ???? ???? ?????
4??? ???? ?? ???
- ??? ????? ?????? ?????? ?? ????? ???? ?????? ?? ?
?????? ??? ??? ?? ???? ??? ???? ? ?? ???? ?????
???? ??? ??? ???? ?? ??? ????? ???? ???? ?? ????. - ????? ?????? ????? ???? ????? ??????? ? ???
?????? ????? ???? ?? ???? ? ?? ?? ???????? ???
?????? ?? ??? ??????? ???? ???? ?? ??? ?? ??
????? ???? ?? ????. - ?? ????? ???? ???? ???? ??? ???? ?????? ???? ??
?? ????? ???? ???? ??? ?? ??? - ???? ????? ???? ?? ??? ????? ????? ???? ???? ???
??? ? ?? ??????? ????? ???? ???? ??? ? ???
?????????
5- ????? ???? ?? ???? ?????
- ????? ???? ?? ??? ?? ???? ????? ????
- ???? ? ??????? ???? ????? ????? ?? ???? ?????
???? - ????? ???? ? ??? ????? ???? ????? ????
6????? ????? ??? ????
- 1- ????? ????? ?? ??? ? ???? ????????? ???? ???
???? ???? ??????? ????? ?? ??? ? ??? ?? ???? ????
???? ?? ??????? ?? ????. - 2- ??????? ???? ?? ????? Na ? Cl ?????? ???
????? ???? ???? ??? ????? ?? ????? ??? ?? ??
??????? ??? ???? ??? ??? ???? ??? ) ?? ????
??????? ??????? ?? ??? ? ?) ?? ???? ?? ?????? ???
??????? ?? ????? ?? ???? ?) ?? ???? ???? ???
?????? ???? ?? ?? ????
73- ?????? ?? ??? ???? ?????
- 1- ????? ??? ????? ?? ??? ? ????? ???? ???? ??
????. ?????? ?? ???? ?? ??? ??? ?????? ???? ????
??? ???? ??? Na ???? ?????? ?? ??? Ca ? K ??? .
?? ??? Mg ? ??? K ?? ???? ?? ???. ??? Cl ? Na
???? ?? ???? ????? ????? ?? ???. - 2- ??? ????? ?? ??? ?????? ? ??????? ??? ???? ??
??? ????? ?? ??? ? ??? ??????? ????? ???? ???? ?
????? ??????? ????. ???? ?????? ?? ?? ???.?????
??????? ?? ??? ?? ???. ???? ???? ?? ???? - 3- ?? ??? ??????? ??? ???? ??? ??? Atpase-H pump
??? ?? ?? ??? ???????? ATP ? ?? ??? ??? ??????
? ???? K ???.
8??????? ????????? ???? ?? ????
- 1-??? ???? ?? ??? ?????
- - ??? ??? ClNa ???? ?? ??? K ? Ca ??? ??? ??
????? ? ?????? ?? ???? ??? ?????? ?? ???. - - ???? ?? ??? ??? ???? ? K ???? ??? ????? ?????
- 2-??? ??? ???????
- - ???? ??? ??? ? ??????? ???? ???????
- - ?? ?????? ????? ???? ??? ?? ??? ???? ?? ???? ?
??? ????? ??????? ???? ?? ???? ?? ????? ????
????? ????? ??? ?????? ????? ??? ???? ????????
????? ?? ??? ??????? ?? ????? ?? ???.
9- 3-??? ???? ?? ????? ??????? ????????? ??
- - ????? ?????? ???? ?? ????????? ?? ???? ?? ????
??? ????? ??? ??? ? ??? ????? ????? ???? ?? ???? - ??? ???? ??????? ??? ???????? ??? ?? ?????.
- 4-??? ???? ?? ????? ???? (???? Pathway)
????????? - ????????? ???? C3 ????? ?? ????? CAM
- ( ????????? ?????? ??????? (Crassulation Acid
Metabolis ????? ??? C3 ???? ???? ??? ?? ???
????? ??? CAM ????? ???? ?? ????. ?? ??? ?????
????? ?? ?? ???? ?? ?? ?????? ?? ?? ???? RDC (
??????? ?? ????? ?????????? ) ??????? ?? ??? ?
?? ??? ?????? ???????? ??????? ????????? ????
?? ???.
10????? ??????????? ?????
- ????? ?? ????
- ????? ?? ????????
- ????? ?? ?????
- ????? ?? ??????? ???
- ????? ?? ?????? ???
115- ??? ???? ??? ????? ??? ?????
- - ?? ??? ???? ??? ????? ?????? ?? ???? ?? ??????
???? ????? ?? ???? ???? - - ?????? ???? ??? ????? ?????? ???? ????? ??????
?? ???? ???. (?? ??? ?? ??? ?? ?????? ????????
?????? ?????? ?? ???? ?? ?????? ?? ?????? ?????
???? ??? ?? ?????? ????? ???? ????) - - ?? ?????? ?????? ???? ?????? ???? ?? ????? ???
?????? ???? ?? ??? ???? ???? ???? ???????
???????? ??? (?? ???? ???? ??????? ???? ????)
?????? ?? ???? ??????? ? ????? ?? ? ?????? ?? ???
ATP ???? ? ?????? ???? ??? ??? ???? K? ????
????? ???? ???.
12- - ????? ???????? ????????? ???? ?? ??? ???? ?? ?
??????? ??? ??? ?? ??? ???? ???? ?? ????? ???
???????? C ????? ?? ????? ????? ??? ???? ????.
?? ??? ???? ?????? ????? ????? ??? ? ???? ??
????? ?? ?????? ? ??????? ??? ????? ????? ?? ???. - - ???? ???????? ????????? ?????? ?? ???? ????
???? ???? ???? ?? ???? ???????? ?? ??? ???? ???
????? ????? ??? ???? ?? ???? ??????? Aldolase
???? ?? ?????.
136-????? ???? ?? ??? ???? ??????? ??
- - ?? ??????? ?? ???? ??????? ?? ??????? ????? ??
???? ???? ?? ??? - -????? ?????? (???? ?????) ?? ???? ?????? ???
???? ?? 100 ????? ?? ???? - - ?? ???? ??????? ???? ??????????????? ? ????
?????? ?? ??? ??????? ??? ????? ? ?? ???? ????
???? ?? ???? ?????? ????? ? ???? ????? ???? ?????
?? ???? ?? ????. - - ?????? ????? ?? ??????? ??? ???? ???? ????? ??
???? ?? ???? ? ???? ???? ?? ??? ????? ?? ?? ???. - - ?????? ?? ???? ???? ???? ???? ??????? ??? ????
?? ???? ???? ?? ???? ????? ???? ?? NH3 ?? ?? ????
???? ???? ?? ???. - - ?????? ?? ?? ??? ???? ???? ?? ????? ????? ?
????? ????? ???. ????? ?? ??? ???? ??? ??????
????? ?? ??? ?? ???? ???????? ?? ??? ??? ? ?? ???
???? ? ???? ?? ????? ??? ?????? ?????.
14 7-????? ??????? ????
- - ?????? ABA ?? ???? ????? ????? ?? ??? ? CK
?????? ???? ?? ????. ??? ???? ?? ???? ?????? ??
??? ?? ????? ?? ????. - - ???? ABA ???? ? Ck ????? ???? ??????? ???? ?
?????? ??????? ????? ?? ????? ???? . - - ???? ????????? ?????? ???? ?? ???? ?? ??? ?
???? ????????? ??? ?? ???? ?? ??? ? ???? ?????
?? ????? ?? ???? ??? ?? ???.
15?????? ? ??? ??? ???
- ?? ???? ?? ???? ???? ??????? ????? ?? ?????? ???
? ??? ??? ???? ??? ????? ????? ??? ????? ??????
?? ?????? ?? ????? ??????? ???? ???? ?????? ??
?????? ????? ???? ? ??? ?????? ??? ???? ??? ????
? ?????? ???? ????? ?? ????.
16- 1-?????? ??? ???? (Halophytes) ??? ?????? ??
?????? ?? ????? ???? ?? ?????? ??? ?? ???? ? ??
??? ???? ?????? ? ??? ???? ??????? ????? ??
????. ??? ?????? ???? ?????? ??? ? ?????? ???
???? ??? (Atriplex) ?? ??????? ??????. -
- 2-?????? ??? ??? ???? (Glycophytes)
- ??? ?????? ????? ????? ??? ?? ???? ???? ????
???? ???????? ????? ?? ?????? ?? ?????????? ??
???? ???? ?????? ?? ????? ????? ?? ???? ?????.
17- ????? ???? ??? ???? ?????
- ??? ????
- ??? ????? ????
- ??? ?????
- ??? ???? ???
18????? ???? ??? ???????? ? ??????? ????
- 1- ???? ??? ????? ???? ??? ??? ???? ?? ?? ???
??? ???? ???? ?? ???? ??? ????? ???? ?? ???? ????
????? ???? ????? ?????? ???? ????. ???????? ????
???? ??? ?? ??? ?? ???. - 2- ???? ??? ????? ???? ???? ????????? ???????
???? ???????
19- 3-????? ???? ?????
- - ????? ???? (Succulence) ????? ?????? ??? ????
??? - - ?? ?????? ??? ??? ???? ???? ?????? ??? ???? ???
???? ??? ??? ?? ?? ?????? ?? ?? ???? ???? ??? ???
??? ?? ??? ????? ????? ?????? ???? ??? ?????? ?
??????? - - ????? ??? ???? ?????? ??? ???? ?? ?????? ?? ???
??? ?? ???? ? ?? ??? ??? ????? ???? ?? ?????.
????????? ?? ????? Succulence ?????? ?????? ???
?? ???.
20- 4- ???? ??? ??????? ??? ?? ????? ?? ??? ?????
?? ?????. - 5- ????? ?? ????? ????? ?? ? ?????? ???? ?? ??
??? ???? ???? ???? ???? ?????? ???. - 6- ????? ????? ?? ?? ???? ?? ???? ??? ?? ???
(???? ??? ??? ???) ??? ???? ????? ????. ???? ???
?? ??? ????? ?? ??????? ???? ????? ?? ????. ????
?? ?????? ?????? ???? ??? ????? ?? ???? ????? ??
?? ???? Pit ?? ???? ???? ???? ?? ??? ? ???? ??
??? ? ???? ???? ?? ?? ???? ???? ?? ????. ?? ???
???? ?? ?? ???? ?? ?? ????? ??? ?? ?????.
21- 7-??? ???? ??? ???? ?? ???? ???? ??? ?? ????
???? ??? ? ???? ?? ??? ? ?? ???? ?? ?? ???? ????
?? ????. ??? ????? ?? ??? ???? ?? ???????? ?????
???? ?? ??? ? ?? ???? ????? ?? ??? ??? ???? ????
??? ???? ?? ???? ?? ???? ??????? ?? ???. - 8- ??????? ?? ????? ????? ??????? ???? ? ????
???? ??? ???? ?????? ??? ????? ???? ???? ?? ????
? ???? ??? ????? ??? ?????? ?????? ??? ???? ???
????? ?????.
22?????????? ?????? ?? ????? ???? (??????)
- 1-?????? ????????
- - ????? ??? ?? ?? ???? ??????? ? ?? ??? ?????
????? ??????? - - ??? ???? ?? ??????? ??? ???? ????? ??? ??? ????
????? ?? ??? Al ??????? ?? ???. - - ?????? ???????? ?? ???? ???? ???? ????? ?? ?? ?
?? ????? ?? ???? ???.
23 ????? ?????????? ?????? ?? ????? ???? 2 - ????
???? ??? (Salt Exclusion)
- ???- ?? ???? ???? ??? ?? ??? ??? ??? ? ???? ????
????? ?? ?????? ?? ?????? ???? ??????? ?? ???. ??
???? ???? ????? ?????? ??? ??? ???? ?? ?? ?????
????? ?? ????? ?? ?????? ?? ???? ??? ?? ????.
24- ?- ????? ?? ???? ??????? ???? ?? ??? ??? ??? ??
?????. ???? ?????? ???? ??? ?????? ?? ????? ?????
?? ??? ?? ???????? ???? ?? ???? ?? ????? ??????
??? ???? ???? Dehydration ?? ??? ????? ??? ???
???????? ?? ??? ????? ?? ??? ????? ??????? ?? ???
?????? ???? ?? ??? ??? ?? ??? ??? ???? ??????
????? ??? ??? ? ?? ????? ??? ?? ????? ??? ??
????. ?????? ???? ??? ?? ??? ??? ? ???????? ?????
?? ??? ????? ??? ????? ??? ???? ? ????? ???? ???.
25- ?- ???? ??? ?? ????? ?? ??? ?? ?? ??? ????? ??
???? ?? ?? ???? ?? ?? ???? ????? (???? ?????)
???? ????. ?????? ???? ??? ??????? ?? ???
?????????? ?? ???.
26 ????? ?????????? ?????? ?? ????? ???? 3 - ????
????
- - ????? ????? ?? ????? ???? ?? ??? ?? ? ???? ???
??????? ?? ???. ?? ????? ?? ???? ?? ???? ??? ????
??????? ?????? ???. - - ???????? ??? ? ??? ????? ????? ???? ????? ???
???????? ????? ??????? ??????? ?? ??? ? ???? ???
????? ?? (??? ?? ??)? ????? ???? ?? ????? ?? ???
? ?? ?? ??? ??? ???? ??? ??? ???????? ????? ??
?? ???? ?????? ?? ?? ??? ? ?? ???? ??????? ?? ???.
27- ???? ??? ????? ?????? ????????? ?? ??? ? ?????
Co2 ?? ?? ?? ??? ??????? ???? ???? ??? ?? ??????
CAM ?? ?? - PEPC
- Co2PEP
Malic Acid - ?? ?????? ?????? ????? ?? ???. ??? ?????? ??????
??? ????? ????? ?? ????? ???? ?? ?? ??? ????? ??
???? ? ?? ??? ?? ??? ????? ?? ????. - Co2
- V MA
- PG (C3)
284-?????? ???
- - ????? ? ????? ??? ?? ????? ???? ? ????? ???
???? ??? ? ??? ?? ??? ????. - - ????? ??? Branching ?? ?????? ?? ???.
295-??????? ??? ?????????
- - ?????? ?????? ?????? ?? ????? ??? ????? ?????
???. - - ??? ????? ? ?????? ???? ? ??? ??????? ???? ??
????? ????? ??? ?? ???? ????? ??? ??????? ???? ??
??? ??????? ???? ?? ?? ????? ???? ???? ?????? ??
???? ???? ?? ???? (???????). ??? ??????? ?
??????? ??? ????? ???? ???? ? ??????? ???????
???? ?? ????. - ???? ??????? ?????? ?? ????????? ?? ABA ????? ??
???. ???? ???? ABA ?? ???? ????? ?? ??? ???????
????? ?? ???. - - ?????? ???? ABA ??? ?? ??????? ?????????
????? ?? ???.
305-????? ??? ??? ???? Salt Glands
- - ??? ??? ?? ?? ?????? ??? ???? ???? ?? ???.???
??? ???? ??? ???? ?? ????? ??? ????? ?? ????? ?
??? ?? ???? ??? ?? ???.
31Role of Salicylic Acid
- It has already been established that a
relatively new plant growth regulator, salicylic
acid (SA) improved the chilling resistance of
maize (Janda et al. 1999) by activating - a new isoenzyme of ascorbate peroxidase,
- a H2O2 scavenging enzyme and by decreasing
ethylene production, -
- accumulation of its precursor, 1-aminocyclopropane
-1-carboxylic acid (ACC) in nonacclimated plants
during chilling (Szalai et al. 2000).
32In order to reveal biochemical and physiological
effects of SA pre-treatment which led to improved
fitness of plants exposed to salt stress.
- Methods
- Tomato plants ( Lycopersicon esculentum Mill. L.
cv. Rio Fuego) were grown in hydroponic culture
both in the presence or in the absence of SA
applied at 10-7-5x10-4 M concentration range. - Six-week-old plants were treated with 100 mM NaCl
and were exposed to salinity stress for a week.
33- . Water potential of plants was estimated using
pressure PMS Instruments and - and osmotic potential was determined in the
expressed cell sap by digital automat osmometer - Moreover, SA increased the rate of photosynthetic
- Chlorophyll fluorescence measurements were made
and ethylene production was determined by GC
(Hewlett-Packard, 5890 Series II). - Total soluble sugar contents were estimated by
the method of Dubois et al. (1956).
34- Results of using SA on salinity resistance
- Long-term incubation of tomato plants in low
concentration of salicylic acid enabled plants to
tolerate salt stress caused - by 100 mM NaCl. Na ions accumulated in the
leaf tissues of treated plants and functioned as
osmolytes without the well-known detrimental
effects of the excess sodium.