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ASCENT OF SAP
B.Sc BIOTECHNOLOGY 08.05.2012 TRANSLOCATION OF WATER
Introduction
Plants absorb water through the roots. This absorbed water reached up to leaves through the stem branches. Most of the water is lost by the transpiration from the leaves. Only small amount of water utilized in vital activities of the plant.
"The upward movement of absorbed water by the under ground roots towards aerial parts of the plant against the gravitation is called Ascent of sap". Process of ascent of sap takes place from a small plant to a quite large plant e.g. in Sequoia semipervense. It has been experimentally proved that water moves upwards through the vessels and tracheids of xylem.
Ringing [girdling] experiment has been done only in dicotyledons because their vascular bundles lies in a ring. While vascular bundles are scattered in monocotyledons, so that this experiment can not be possible.
Structures involved in Ascent of Sap: Various experiments like girdling, staining and plugging, indicate that the xylem tissue is mainly responsible for the movement of water. As xylem consists of tracheae otherwise called vessels, they form a system of fine channels running from roots to all other regions of the plant body and form a beautifully branched supply system, which is almost similar to that of arteries in animals.
Many theories have been put forwards by various co-workers to explain the mechanism of ascent of sap. (1) Vital Force Theory (2) Root Pressure Theory (3) Physical Force Theory
J.C. Bose measured this pulsation activity by "Electric probe" (Galvanometer) apparatus Molish A scientist named Molish who supported the pulsation theory of J.C. Bose and he introduced a detail description of pulsation theory. According to him the pulsatory activity increased to 14 seconds by the application of narcotics drugs on the plant.
Objections
The view of vital theory discarded by Straburger and Boucheries. They proved by their experiment that the living cells are not essential for ascent of sap. Because when the living cells are killed (destroyed) by picric acid or HgCl2 solution even then ascent of sap continues.
The maximum value of root pressure falls in the range of 2 - 3 atmosphere. The liquid which flows from the freshly decapitated stem of the plant is called bleeding. This is also due to root pressure. Guttation is also takes place due to root pressure. Ascent of sap through the root pressures only possible in small and herbaceous plants. Therefore it is only applicable for small and herbaceous plants.
The process of ascent of sap can not be explained by root pressure because it has following limitations (1) Gymnosperms lack root pressure and the conifers are very tall plants. (2) The maximum transpiration is found during the summer and plants require more water but in that time root pressure is found to be very less. (3) The rate of ascent of sap is found rapidly even in the absence of root pressure. (4) The rising up of water can not be possible at 2 atmosphere in tall plants. Upto 200 feet height plants require 10 atmosphere root pressure to push the water to the top of the plants. (5) A negative root pressure is found in some of the desert plants. (6) Root pressure is found less in summer and more in winter. (7) Root pressure takes place in the special conditions when the rate of absorption is higher and transpiration rate is minimum. Hence, root pressure is not significant in most of the plants. It is only effective in low transpiring herbaceous plants. Therefore, theory explain partially the mechanism of ascent of sap.
Limitation of this theory as follows : (a) Xylem vessels are not freely immersed into the water. (b) The ends of vessels and tracheids are closed while ends of capillaries are open. (c) The diameter of the capillaries should be very thin because capillary with narrow lumen encourages the entry of more water. But in tall plants the diameter of vessels and tracheids is more. At this height capillary force does not operate. (d) Tall plants like Gymnosperms having tracheids instead of vessels which have many transverse septum. Therefore, tracheids are homologous as capillary. But still ascent of sap occurs. (e) Lifting power of capillary is not much and can not account for the rise of water exceeding 3 meters. It is capable for few centimeters rise only. Capillary theory is completely discarded on the basis of above reasons.
The process of ascent of sap a continues through the medium of above mechanisms. Evidences in support of Dixon's theory : (1) Normally, the rate of absorption is equal to the rate of transpiration water does not come out from the cut end of the stem during transpiration. When this cut end is watered, it absorb water inside the stem. This justifies the water tension in the stem. (2) McDougal is found that daily contraction in stem of the trees with the help dendrograph apparatus. Maximum contraction is found during a maximum transpiration. (3) The maximum value of osmotic pressure of the cells of leaves is found during noon. Because the amount of water is minimum in that time
Although air bubbles are present in the xylem, but they are unable to move much distance because (i) xylem vessels do not continue for long distances, pits are present between them. (ii) Tracheid are also present along the vessels as alternative path. Pits are also present in them. (iii) Membrane and wall is present on the pits which is permeable. to water, but air bubbles do not pass out through them. Air bubbles may dissolve in water. Root pressure also helps to remove air bubbles.
The rate of transpiration is higher than that of rate of ascent of sap during the day time and is called absorption lag. Russel and Wooley found the ratio 50 : 1 by comparative study of water movement in apoplast and symplast. The origin of root pressure is an active process. Factors affecting Ascent of sap (1) Amount of water in soil. (2) High temperature (3) High atmospheric humidity (4) High atmospheric pressure (5) Wind velocity (6) Number of stomata in leaves
However anatomical studies have failed to show any pumping action in the Xylem.Further numerous experiments have demonstrated clearly that living cells are not involved in the translocation of sap in the stem.
ROOT PRESSURE THEORY Acc to Root pressure theory,the positive pressure developed in the roots pumps the sap up in the xyulem ducts.This theory seems to be applcable to many species,which generate considerable amount of root pressure.For example,Grapevines have been shown to generate root pressure up to 5-6 bars,which can support a water column upto 150-160 ft
However, root pressure cannot account for sap translocation in many situations.In many species,root pressure has not been observed at all and in many others,it is low that it cannot account for upward translocation even upto a few ft.
COHESION THEORY BY IRISH BOTANIST HENRI DIXON and a Physicist Jon Joley(1894) Supported by E.Askenasy(1895)and Renner(1895).
Water has a high Cohesive( Mutual attraction) force.This is primarily because of the formation of Hydrogen bonds between water molecules.Water also shows forces of adhesive with the walls of xylem vessels and tracheids.Because of Cohesive and adhesive forces,it exists as a column in the xylem elements.
Organismal Circulation
Unicellular Organisms Autotrophic Multicellular Organisms (Heterotrophic Multicellular Organisms)
http://botit.botany.wisc.edu/courses/img/Botany_1 30/Movies/Slime_mold.mov
The correct taxonomic affiliation is unclear. It has been treated as Fungus and Protist. Further study is needed to resolve its position.
http://www.microscopy-uk.org.uk/mag/imgnov00/cycloa3i.avi
The shoot organ system is photoautotrophic, taking in CO2 and releasing O2 in daylight.
Diffusion is sufficient to exchange gases. But solutes need to be circulated in the large plant body as diffusion is too slow!!
Stem
Lateral roots
The root organ system is chemoheterotrophic, taking in O2 and releasing CO2 in the darkness of the soil environment.
The shoot system produces carbohydrates (etc.) by photosynthesis. These solutes are transported to the roots in the phloem tissue:
Branch
Translocation
Stem Transpiration
The root system removes water and minerals from the soil environment. These solutes are transported to the shoot in the xylem tissue:
Translocation
Lateral roots
Transpiration
Taproot
Node Internode
Because these pathways involve solutes in water passing in the adjacent tissues of a narrow vascular bundle, this is a circulation system!
Transpiration and Translocation The water is moving up the xylem, and down the phloem, making a full circuit!
Stem Transpiration
Translocation
Lateral roots
Taproot
Epidermis
Cortex Pith
Ground tissue
Vascular bundles
vascular bundles
As a monocot plant grows in diameter, new bundles are added toward the outside for increased circulation to the larger plant body.
Is this slice from a young or a mature part of the corn stem? Lets take a closer look at the vascular tissues
Translocation
Transpiration
phloem fibers
Support of Stem functional phloem Translocation vascular cambium Cell Divison: More Xylem and Phloem
xylem
Transpiration
Each year the vascular cambium make a new layer of secondary xylem and secondary phloem
Dicot stem anatomy: four year-old stem (3 annual growth rings) phloem etc. = bark All of these tissues were added by the vascular cambium!
xylem = wood
cambium phloem
heartwood
pith
When flowering plants are young, water needs are limited, tracheids suffice. The walls are strengthened with secondary thickenings including lignin. Protoxylem have stretchable annular or helical thickenings.
As flowering plants age and grow, water needs increase, and tracheids need to be supplemented.
Flowering plants evolved xylem cells with larger cell diameter and perforated end walls to increase water flow. Vessels have perforated end walls or lack end walls, but lateral flow between cells is still through pits.
The huge vessel transports lots of water longitudinally, and shows lots of pits for lateral transport
The huge vessel transports lots of water longitudinally, and shows lots of pits for lateral transport
Vessels, Tracheids have different distribution patterns. Some produce big vessels only in spring wood Others produce vessels year-round.
This sketch is showing the importance of lateral transport. In both transpiration and translocation materials must move radially to the interior and to the exterior as well as up and down the plant. O2 in and CO2 out
Dicot stem anatomy: 2-year old stem showing ray and periderm
phloem
1996 Norton Presentation Maker, W. W. Norton & Company
Rays transport sugar from the phloem toward the interior to keep pith and xylem parenchyma fueled. Rays transport water and minerals from the xylem to the exterior to keep the periderm, cortex, and phloem parenchyma hydrated.
Xylem and Phloem: tissues with many cell types but conduction function toward pith
radial transport
toward cortex
112 m tall (367.5 feet)! This tree is more than ten times taller than is theoretically possible based solely upon the length of the column of uncavitated water. How could this be achieved?
http://www.nearctica.com/trees/conifer/tsuga/Ssemp10.jpg
Transpiration: root pressure (osmotic push) Solutes from translocation of sugars accumulate in roots.
1996 Norton Presentation Maker, W. W. Norton & Company
Water from the soil moves in by osmosis. Accumulating water in the root rises in the xylem. Water escapes from hydathodes.
guttation
Transpiration: root pressure (osmotic push) The veins (coarse and fine) show that no cell in a leaf is far from xylem and phloem (i.e.water and food!). The xylem of the veins leaks at the leaf margin in a modified stoma called the hydathode. These droplets are xylem sap.
http://img.fotocommunity.com/photos/8489473.jpg
Capillarity: maximum height of unbroken water column glass tube vacuum created gravity pulls water down
The small diameter of vessels and tracheids and the surface tension of water provide capillary (climb).
10.4m
Cohesion of water, caused by hydrogen bonds, helps avoid cavitation. A tree taller than 10.4 m would need some adaptations to avoid cavitation
water
Conifer stem anatomy: pine xylem tracheids with pits, xylem rays
vascular cambium
In spite of the limitations of tracheidsonly xylem, conifers are among the tallest of trees!
ray parenchyma
P low
P high
These pit features allow conifers to be very tall and still avoid cavitation in their xylem cells. As pressures change between adjacent cells, the torus movement blocks catastrophic flow that would result in cavitation.
water Water evaporating from a porous clay cap also lifts the mercury! mercury
Transpiration can lift the vacuum mercury above its normal cavitation height!
76 cm
mercury
Leaf
High Pressure
plasmodesmata
Low Pressure
Modified from: 1996 Norton Presentation Maker, W. W. Norton & Company
Low Pressure
High Pressure
Transpiration Evaporation: Water evaporates from mesophyll into atmosphere. Water molecules are pulled up the xylem by virtue of cohesion. Capillarity: Water climbs in the xylem cell walls by adhesion. Water molecules follow by cohesion. Root Pressure: Water moves into the root because of solutes from phloem. Pressure pushes the water up the stem. Node Internode Node Leaves
Branch
Stem Transpiration
Translocation
Lateral roots
Taproot
Figure 36-3 Page 793
Translocation
Branch
Leaf = Source Photosynthesis produces solutes. Solutes loaded into phloem by active transport. Water follows by osmosis, increasing pressure.
Root (etc.) = Sinks Solutes removed from phloem by active transport. Water follows by osmosis, reducing pressure. Pressure = Bulk Flow The pressure gradient forces phloem sap away from leaves to all sinks (bidirectionally).
Stem Transpiration
Translocation
Lateral roots
Taproot
Figure 36-3 Page 793
Thank you!
Next Lecture: TRANSPIRATION