Title: Transport system
1100311 ?????????????
11
- ??????????????????
- ????????????????????????????????????????????????
???????? ????????????????????????????????
?????????????????? - ?????????????????????????
- ???????????????
- ???????????? ??????? ??? ?????????
??????????????????
2100311 ?????????????
?????????????????????????? ??????????????????????
?????????????
- Transport system
- Photosynthesis
- Respiration
- N and S metabolism
- Development
3100311 ?????????????
- ???? Physiology for Crop Production ??????????
- ?????????????????????????????????????????????
- ???????????????????????????????????????????
????????????????????????????????????????
- ??????????????????????????
- ?????????????????????????????????????????????????
???? ?????????????????????????? - ?????????????????????????????????????????????????
??? - ??????????????????????????????????
??????????????????????????????
4100311 ????????????? ????????????????????????????
???
? ????????????????????????????????????????????????
? ?????????????????? xylem ? ??????????????????
phloem
5100311 ???????????????????????????????
??????????????????????????????
- ???????????????????? ??????????????????????
??????????????? - ??????????????????????
- ????????????????????????????????????
??????????????
- ?????????????????????????????????????????? ?
?????? - ??????? (diffusion)
- Mass flow
6100311 ???????????????????????????????
??????????????????????????????
Diffusion
????????????????????????? ????????????????????????
??????????????????? ??????? ?????????????????????
???? ????????????????????
- ??????????????? (J) ??????????
- ????????????????????????? (DC)
???????????????????????????? (Dx) - ?????????????????? (k)
Ficks Law
???????? ???????? ?????????????????????????????
7100311 ???????????????????????????????
??????????????????????????????
????????
- ?????????????? i ?????? a Ca 20 moles m-3
?????? b Cb 10 moles m-3 - ??????????????? a b ??????? 0.05 m
- ???????????????????????????? 0.01610-3 m2 s-1
???????? Fick
?????????????????? i ??? a ?? b ??????? 0.0032
moles m-2 s-1
8100311 ???????????????????????????????
??????????????????????????????
Mass flow
????????????? mass flow ??? ??????????????????????
?????????????? ??? ??????? ???????????????????
????????????????
- ???????? mass flow (Jv) ??????????
- ????????????????????? ( D P) ???
????????????????????????? ( Dx) - ??????????????????? (hydraulic conductivity, Lp)
Darcy Law
???????? ???????? ?????????????????????????????
9100311 ???????????????????????????????
??????????????????????????????
????????
- ????????????? a Pa 0.4 MPa ?????? b Pb 0.2
MPa - ??????????????? a b ??????? 10 m
- ????????????????????????????? 0.05 m2 MPa-1 s-1
???????? Darcy
???????????????????????????????? a ?? b ???????
0.001 m s-1
10100311 ???????????????????????????????
??????????????????????????????
??????????????????????????????????????????????
?????????????????????? ???
Water sugar H.pressure
Water sugar H.pressure
pure water L.pressure
pure water L.pressure
Semi-permeable membrane
Semi-permeable membrane
?????????????????????????????????????????????????
??????????????????????????
11100311 ???????????????????????????????
??????????????????????????????
???????????? Free energy ??? ????????????/???
Chemical potential (m)
- ????????????
- ???????????????????????????????????????
- ???????????????????????????? ????????????????????
??????????? ?????????????????????? - ????????????????????????????????????????
?????????????????????????????????? - ???????????????? extensive
- Chemical potential (m)
- ??????????????????
- ???????????????? intensive ??????????????????????
??? ?????????????????????????????? free energy
12100311 ???????????????????????????????
??????????????????????????????
????
- ????????? ?????????????????????? Diffusion ????
Mass Flow ??????????????????????????????????????
???
Diffusion Fick Law
Mass flow Darcy Law
- ????? ???????????????????????????????????????????
??????????????????? ?????????????? ???
???????????????????????????????????????? chemical
potential
13100311 ???????????????????????????????
??????????????????????????????
Additive property
?????????????????????????????????????? additive
??????? chemical potential ????? ???????????????
chemical potential ???????????????????????? ????
??????????? ??????? ?????????? ??????????????? ???
14100311 ???????????????????????????????
??????????????????????????????
????????????? chemical potential ?????????????????
1 Chemical potential ?????????????????????????
(osmotic potential)
R gas constant, 8.3 J mol-1 Ko , T Kelvin
Ko , a activity of i
?????????????? g activity coefficient 1,
For solute ai g C
(C, molality)
For solvent, ai g N,
(N, mole fraction)
15100311 ???????????????????????????????
??????????????????????????????
2 Chemical potential ???????????????????
(pressure potential)
V partial molal volume, m3 mol-1 P
Pressure, J m-3 (Pa)
3 Chemical potential ???????????????????????????
(gravitational potential)
m mass/mole, kg mol-1, g 9.8 m s-2,
h height, m
4 Chemical potential ??????????????????????
(electrical potential)
Z valency, F Faraday constant (96487
coulombs mol-1), E volt
16100311 ???????????????????????????????
??????????????????????????????
Chemical potential ??????????????????????????????
???????????????????????? additive ??????????
??????????? chemical potential ??????????????????
????????????????????????????????????? chemical
potential ????????
J mol m-2 s-1 k coefficient, mol2 J-1 s-1 m-1
17100311 ???????????????????????????????
??????????????????????????????
Example
T293 K
Cell
Nw 0.990 P atm Lp 22.8410-4 mol2
J-1 m-2 s-1
Nw 0.995 P atm
Chemical potential
J mol-1
Flux of water into the cell
moles m-2 s-1
At equilibrium
18100311 ???????????????????????????????
??????????????????????????????
????
General forms are
Diffusion Fick Law
Mass flow Darcy Law
Multiple forces
????
At equilibruim
Dm 0
Ji 0
19100311 ???????????????????????????????
??????????????????????????????
?????????????????????????????????
- ???????????????????????????????
????????????
???????????? chemical potential ????????? A
?????????? C ??????? ????????????????????????????
??????????????????
chemical potential ?????????? A ??????? ??? C
- ????????????????????????????????????????
chemical potential ??????????????? ????
????????????????????????????????????
??????????????????????????????????????????
20100311 ???????????????????????????????
??????????????????????????????
??????????????????????
1. ??????? solvent, N 1 ?????????? solute, C1
molality 2. ???????? (T) 25 oC ???? 298 oK 3.
??????? (P) ?????????????????? 4. ???????
??????????????????????? 5. ?????????? (E) 0
21100311 ???????????????????????????????
??????????????????????????????
???????
Water potential (Y).
????????????????????? chemical potential
?????????????????????????????? ??? chemical
potential ????????????????????????????????????????
?????????? ???????
????????????????????????????
22100311 ???????????????????????????????
??????????????????????????????
???????
- ????????? ???????????????? perfect semipermeable
s reflection coeeficient ?????????????????????????
?????????????????? ???????????????????????????????
? 1 ??????? 100 0?????????????
- ??????????????? colloids ???? proteins????? ????
?????? clay ?????? - ????? molecules ??????????????????????????
matric potential yt
(ys ???? yt lt 0)
23100311 ???????????????????????????????
??????????????????????????????
???????????????
- ????????????? ????????????????????????????????
????????????????????????????????????? partial
pressure (e) - Total pressure partial pressure of gas1
partial pressure of gas2 ...gasn - Partial pressure ???????????????????????
??????????????????????? saturation vapour
pressure (es)
24100311 ???????????????????????????????
??????????????????????????????
vapour
??????????
Liquid
???????? Raoult
?????? Nw/Nw ?????????? 1 ???
25100311 ???????????????????????????????
??????????????????????????????
???????????????????????????????????? (xylem) ???
????? (phloem)
???????????????????? ????? ?????
??? ?????? ???????
Phloem Transport
??????????????
Xylem Transport
??? ???????
?????????????????????
???
26100311 ????????????? xylem transport
?????????????????? xylem ??????????? 3 ???? 1
???????????????? xylem ?????? 2 ????????????????
xylem ??????????????? 3 ??????????????????? xylem
????????????????????????????????? (?????????)
27100311 ????????????? xylem transport
28100311 ????????????? xylem transport
29100311 ????????????? xylem transport
????????????????
Epidermis
Endodermis and casparian strip
Cortex
xylem
Phloem
Molecule
Pericycle
- ???? membrane ????????? 2 ?????
Water Ions
membrane epidermis cortex endodermis
Xylem
membrane endodermis pericycle
Symplasm
Apoplasm
Apoplasm
30100311 ????????????? xylem transport
???????????? membrane
Hydrophobic (Nonpolar)
Hydrophillic (Polar)
- Proteins ?????? amino acids ?????????????
peptide bond
31100311 ????????????? xylem transport
- ???????????????????????? Fluid mosaic model
Integral protein
Peripheral protein
Bilayers phospholips
- ???????????????? nonpolar ????????????
hydrophobic interactions - polar ???????????????????? ???????????????????
???? H bond - ???????????????????????????????
??????????????????????????????????????????????????
- ?????? proteinlipid 5050 ??? mitochondria ????
chloroplast ????????? 8020
32100311 ????????????? xylem transport
33100311 ????????????? xylem transport
????????????????? membrane ????????????????????
Water dipolar small size
Ions polarity larger size
Aquaporin
Carriers / channels
34100311 ????????????? xylem transport
?????????????????????????????????? membrane
????????? proteins
Physical
1. Saturation kinetics
Enzymic reaction
??????????????????
D ???????????
???????????????????????? membrane
2. Protein inhibitor
Eg. 2,4 dinitrofluorobenzene, cyanide
3. Selectivity NKP gt others with the same Dm
???? ????????? proteins ?????????????
phospholipids ???????
35100311 ????????????? xylem transport
??????????????????????????????????????????
membrane
1. ?????? Dm
Passive transport
High m
Low m
??????????????????????????????????????????????
passive
2 ??????????????????????
Active transport
High m
Low m
?????????????????????????????? ??????? active
36100311 ????????????? xylem transport
- Water passive transport, channel aquaporins
37100311 ????????????? xylem transport
Passive transport of ions channel and carrier
proteins
38Primary active transport-Pumps
100311 ????????????? xylem transport
39100311 ????????????? xylem transport
Primary active transport
40100311 ????????????? xylem transport
Secondary active transport
41100311 ????????????? xylem transport
???????????????????? active
Ion-pump and carrier hypothesis
ATP
ADPPi
H-ATPase
H-ATPase
OH-
HOH
HOH
H
Carrier
_
Carrier
_
_
_
membrane
membrane
Pmf D Electric DH
Metabolic energy (ATP)
42100311 ????????????? xylem transport
ADPPi
ADPPi
HOH
OH-
H
Carrier1
Carrier2
membrane
membrane
D Concentration
Symport (H and ion)
43100311 ????????????? xylem transport
????????????????? active ???? passive
???????? R 8.3 J/(mol K) F96487 C/V
??????????? mM ?????? ????? Na 1.0 14.0 K 0.
1 119.0 Cl- 1.3 65.0
?????????????????? Eo-Ei 0.138V ????????
T293 K
44100311 ????????????? xylem transport
Nernst equation
?????????????
45100311 ????????????? xylem transport
1 ???????????????? xylem 2 ???????????????? xylem
??????????????? 3 ??????????????????? xylem ???
?????????????????????????????? (?????????)
46????????????? xylem
100311 ????????????? xylem transport
47100311 ????????????? xylem transport
???????????????? xylem ???????????????
1. ????????????? root pressure 2. Capillary 3.
??????????????????????? Transpirational pull
48100311 ????????????? xylem transport
????????????? root pressure
Endodermis
Active transport of ions
?????????????? ions inside gt outside Mole
Fraction (N) ?????? outside gt inside
passive transport of water
Endodermis
??????????????????????
Endodermis
???????????????????? mass flow ?????????????
xylem ???????????????????????????
49100311 ????????????? xylem transport
???????????????? Capillary
- Capillary ????????
- Adhesion ????????????????????????????????
- Surface tension ?????????
- Cohesion ????????????????????????????
Adhesion Surface tension Cohesion
Adhesion Surface tension
No force
Only adhesion
50100311 ????????????? xylem transport
G cosa G
Surface tension (G )
a
Tension
a
W
??? a ????? ?????? a 90 ?????? 0 ?????? a
0 ????????????
??????????? Young and Du Pre
????? Ad Co, cosa 1, a 0, ?.?
???-????????????????? ???? ??????? ???? cell
walll ???????? Ad 1/2Co, cosa 0, a 90, ?.?
???-?????????????????????????? ??? polyethylene
51100311 ????????????? xylem transport
G cosa ( 2p r)
????????????? ??????????????????????
(p r2 h) r g
????????
Xylem radius (r) 2010-6 m, G 0.0728 N/m, a
0, r 1000 kg m-3
52100311 ????????????? xylem transport
??????????????????????? transpirational pull
??
?????????
??? xylem
??
??????????????? cell wall ????????????????
?.? ????? , P - 0.8 MPa ??????, P -
0.2 MPa
??? xylem
?????????
53100311 ????????????? xylem transport
???????????????????????????? xylem
- ??????????????????????? xylem ??????? mass flow
??????????????????????? - ????????????????????????????? Darcy
????
?????? s ??? reflection coefficient
??????????????? 0 ?????????????
h ?????????????????????????????? r ????? ??? l
??????????
54100311 ????????????? xylem transport
????????
??
P -0.8 MPa (MJ/m3)
l 10 m, r 2010-6m, h 10-9 MPa-s
???
P -0.5 MPa
(-0.8)-(-0.5) -0.3 MPa
(1810-3 kg/mol) 9.8 (m/s2) (10m) -105
J m-3 0.1 MPa (1810-6 m3/mol)
0.001 m s-1
55100311 ????????????? xylem transport
????
- ??????????????? xylem ???? ??? mass flow
??????????????????????????????????? - Root pressure ??? capillary ?????????????????????
????????????????? xylem ????????????????????????
?????????????? ??????????????????????????????????
xylem
56100311 ????????????? xylem transport
???????????????????????????????
?????????????????? xylem ??????????? 3 ???? 1
???????????????? xylem ?????? 2 ????????????????
xylem ??????????????? 3 ??????????????????? xylem
??? ?????????????????????????????? (?????????)
57100311 ????????????? xylem transport
58100311 ????????????? xylem transport
?????????
??????????????????????????????????
??????????????????????????????????????? Fick
???????????????????????????????????
????? C mol m-3, R s m-1
59100311 ????????????? xylem transport
1 ??????????????????????????????????
D Cwv
??? Gas law
?????????
??? ew ??? P (partial pressure ???????????????)
??????
R ???????????? ??? T ????????
60100311 ????????????? xylem transport
??????????????????????
ew
1.1 ?????????????????????? ??? ????????
1.2 ??????????????????????????????? ??? ????????
????????????????????????? ? ???????? T
T ( K), es (Pa)
61100311 ????????????? xylem transport
2 ????????????????????????????????
Rl
???????????????????? boundary layer
Rb
Uniform air, Velocity and Cwv equal
??
Larminar and turbulent layers, D velocity and DCwv
?????
???????????????????????????????????????? larminar
??????????????????? db
??????
C 0.004 m s-1/2
62100311 ????????????? xylem transport
63100311 ????????????? xylem transport
?????????????????????????
Rst
1. ??????????????? (dst) 2. ?????????????
(rst) 3. ??????????????????? (asa) 4.
?????????????? ??? ?.?. ?? (n)
rst
dst
a
64100311 ????????????? xylem transport
Monocots
65100311 ????????????? xylem transport
??????????????????????????????????????????
Rias
A/Aias
?????????? ?.?.?? ??? ?.?. ???????????
66100311 ????????????? xylem transport
67100311 ????????????? xylem transport
??????????????
??????????????????????????????????? Cwv ???
68100311 ????????????? xylem transport
???????????????????????????????????? 298 K
??? Cwv ????????????
??? Cwv ??????????????????
69100311 ????????????? xylem transport
????????????????? ????????????????????????????????
??
Dw at 25o0.2510-4 m2 s-1
l 0.15 cm, V 1.50 m s-1
stomatal area/leaf area
Leaf area / intercellular area A/Aias 2
70100311 ????????????? xylem transport
??????????
- ?????????????????????? ??????????????????????????
???????????????? - ???????????????????????? ???????????????????????
? - ??????????????????????????????? ???????????? 20
????????????????? - ?????????????????? ???????????? 80
?????????????????????????????????? - ???????? Penman ?????????????????? 2
???????????????????????????
71100311 ????????????? xylem transport
72100311 ????????????? xylem transport
??????????????????????????????????????????????????
?????????
?????????????
Ysoil ??????????? lt--- ?? ??????????
????????????????????? ???? ????????????? ???Yair
???????????????? lt- ??????????????????????????
???? ???????????
Rrt ??????????????? endodermis (casparian
strip) ????????????? passage cells Rxylem lt- 8hl/
r2 lt- ?????????????????? xylem Rleaf lt-
??????? intercellular air space (dias, A/Aias)
??????????????????????? Rst lt-
?????????????????? (dstrst) Rb lt-
???????????????????????????
73100311 ????????????? xylem transport
?????????????????
Jion lt- Jv ????????????? xylem ??????? mass flow,
???????????????????????????????????????? lt-
Active transport lt- ATP lt- O2 ?????????? glucose
lt- ????????????? phloem ??????????????????? lt-
???????????? membrane lt- pH
74100311 ??????????????????????????????? Phloem
Transport
????????????????????????????? source
2. Phloem transport
3. Starch
Sucrose
3. Vacuole
CO2
ATP
G 1 P
G-1-P/ F-6-P
Gl/Fr
Calvin cycle
Triose P sugars (DHAP GAP)
Triose Psugars
Pi
Chloroplast
1.Glycolysis
(Eg. Soybean stores starch in chloroplast but
maize stores sucrose in vacuoles)
75100311 ??????????????????????????????? Phloem
Transport
????????????? Carbohydrates ?? chloroplast
Starch
Specific to Starch
CO2
G 1 P
Calvin cycle
Chloroplast
Common pathway
Triose P sugars
Pi
??????? ?????????????????????????????????
????????????????????????????? ???????
?????????????????????? ???????????????????????????
????????????????? ???????????????????
76100311 ??????????????????????????????? Phloem
Transport
?????????????????
77100311 ??????????????????????????????? Phloem
Transport
78100311 ??????????????????????????????? Phloem
Transport
79100311 ??????????????????????????????? Phloem
Transport
Amylose single chain of gt 10,000 glucose linked
by 1, 4 glycosidic linkage
Amylopectin, branched chains of glucose
1, 6 glycosidic linkage by Branching enzyme (Q
enzyme)
80100311 ??????????????????????????????? Phloem
Transport
???????????????
Glucano hydrolase
Alpha- amylase
Amylose
Dextrins
Amylopectin
Beta- amylase
81100311 ??????????????????????????????? Phloem
Transport
G 6 P
G-1-P
82100311 ??????????????????????????????? Phloem
Transport
????????????????????????????????????????????
G-1-P
G 6 P
F 6 P
P
ATP
Fructose bisphosphatase
Phosphofructokinase
F 1, 6 BP
High energy state ???????????????
Low energy state
3PGal DHAP
83100311 ??????????????????????????????? Phloem
Transport
At sources (fully mature leaves)
1. Oxidation (Respiration) 2. Sucrose
phloem transport to sink 3. Temporaly reserves
(Starch in chloroplast)
Triose sugars 3PGal DHAP
Photosynthesis
CO2
84100311 ??????????????????????????????? Phloem
Transport
??????????????????? sucrose
2 Pi
UTP
P-P UDP
2 (3PGal DHAP)
G-1-P/ F-6P
Sucrose
Common pathway
Specific pathway
85100311 ??????????????????????????????? Phloem
Transport
Common pathway
86100311 ??????????????????????????????? Phloem
Transport
87100311 ??????????????????????????????? Phloem
Transport
88100311 ??????????????????????????????? Phloem
Transport
89100311 ??????????????????????????????? Phloem
Transport
???? ????????????? di ??? polysaccharides
other sugar or Primer
NTP
P-P
NDP
Pi
Disaccharides or polysaccharides
DHAP 3 PGal
G 1-P
NDP-G
90100311 ??????????????????????????????? Phloem
Transport
ATP
ADPPi
Phloem transport
ATPase
Apolasm
H
Symport
Sucrose
Symplasm
Sucrose
Sieve element and companion cell complex
Source cell
Intermediate cell
Phloem
Trapping model Sucrosegalactose raffinose
- Selective and saturation kinetic
- sucrose 80 of phloem organic content
- Active 1-2mM500-1000mM sucrose
91(No Transcript)
92100311 ??????????????????????????????? Phloem
Transport
Pressure flow
Sucrose
Sucrose
Source
Sink
1. Active transport ?????????????????????
?
Active or passive transport ???????????????????
????
2. ???????????????????????? Phloem
?????????????????
??????????????????????? Phloem ???????????????????
3. ???????????????????????????? source ??? sink
???????????????? phloem ????????????? mass
flow ??? source ????? sink
93100311 ??????????????????????????????? Phloem
Transport
????
- Phloem loading
- Apolast active transport
- symplast intermediate cell (trapping model)
- Phloem unloading
- Symplastic route meristematic tissue
- ????? apolast
- ??????????????? sucrose ????? vacuole ???
parenchyma ??????????? sucrose ??????????????
??????????????????? sucrose ????????? sink ????
??? unload ?????????????????? passive
94100311 ??????????????????????????????? Phloem
Transport
????????????? ?????? sucrose ??? sink
95100311 ??????????????????????????????? Phloem
Transport
96100311 ??????????????????????????????? Phloem
Transport
By invertase
Sucrose
Glucose Fructose
- ?????????????????????
- ??????????? sucrose ??? phloem ???? ??????
sucrose ???????????????? phloem ??? passive ??? - ????????????????????????????
By sucrose synthetase
Sucrose
G1 P F1-P
G1-P UTP Fructose
- ????????????????????????
- ?????????????? sucrose ??? phloem ???
???????????????????? sucrose ????????????????????
active - ???????????????????????????
97100311 ??????????????????????????????? Phloem
Transport
Further change of transported carbon
1. Oxidation (respiration)
2. Synthesis of polysaccharides, eg. cellulose,
starch, etc
98100311 ??????????????????????????????? Phloem
Transport
Source
Sink
3. Starch
Phloem transport
Triose P
Glucose Fructose
2. Sucrose
Sucrose
1. Oxidation
1. Oxidation
Intermediates (C-skeletons)
ATP/NADH/NADPH
ATP/NADH/NADPH
Turn over of Proteins Lipids DNA RNA
Proteins Lipids DNA RNA
Polysaccharides Starch
Cellulose Hemicellulose Pectins
99100311 ??????????????????????????????? Phloem
Transport
????????????? cellulose
100(No Transcript)
101100311 ??????????????????????????????? Phloem
Transport
102(No Transcript)
103100311 ??????????????????????????????? Phloem
Transport
??????????? cellulose
H2O
b Glucose
Cellulose
Cellulase cellubioase
Pectin
Primer
Polymer of Galacturonic acids
UDP-Galacturonic acid
a 1-4 glycosidic linkage
UDP
104100311 ??????????????????????????????? Phloem
Transport
Hemicellulose xylans, mannans, galactan
Xylans Polymer of xylose, b 1-4 glycosidic
linkage
Primer
UDP-xylose
Polymer of xylose
UDP
- Mannans
-
- Polymer of Manose glucose galactose, b 1-4
glycosidic linkage - Ration ManoseGlucosegalactose 311
- Galactan
- Polymer of galactose, 1-4 glycosidic linkage
- Side chain disaccharide of Galactose-Arabose
105100311 ??????????????????????????????? Phloem
Transport
Primer NDP
NTP P-P
Monophosphate sugar
NDP- monosaccharide
Polymer of monosaccharides
Cell wall Cellulose hemicellulose pectin
lignin glycoprotein
Wall expansion
Auxin
membrance-bound enzyme
Efflux of H
Break of pectin
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