LEC 3.

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.

1 . The transverse section of this stem confirms because only vessels and tracheids are stained with eosine dye. So that it is proved that the ascent of sap take place through the xylem .Balsam • Take aqueous solution of eosine in a beaker and immerse the cut end of (stock) of balsam plant in it. After sometimes strips of red colour are seen in the stem.Experiment No.

Malpighi and Stephen Hales. .Ringing Experiment • First of all ringing experiment was carried by Hartig. This experiment makes it clear that water moves up only through vessels and tracheids of the xylem. Now placed this experiment in a suitable environment for 2 .3 days.Experiment No. 2 . • We take a branch with leaves immersed in beaker which is filled with water. Now remove all the tissues which are present above the xylem [cortex. Then we observe that leaves did not wilt. cambium and phloem].

• Ringing [girdling] experiment has been done only in dicotyledons because their vascular bundles lies in a ring. . so that this experiment can not be possible. While vascular bundles are scattered in monocotyledons.

• Structures involved in Ascent of Sap: Various experiments like girdling. . As xylem consists of tracheae otherwise called vessels. which is almost similar to that of arteries in animals. indicate that the xylem tissue is mainly responsible for the movement of water. 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. staining and plugging.

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• (1) Vital Force Theory (2) Root Pressure Theory (3) Physical Force Theory .Theories of Ascent of sap • Many theories have been put forwards by various co-workers to explain the mechanism of ascent of sap.

The opinion of the various scientist for ascent of sap as follows : .(1) Vital force theory • Vital theories presumes that ascent of sap is due to the vital activity of living cells of the plants.

(A) Westermaier's theory • Westermaier was the first man who proposed his view about the vital force theory. He explained that force for upward conduction of water is provided by the xylem parenchyma cells and vessels and tracheids are simply acting as water reservoirs. .

A pumping action starts as a result of this changes that causes ascent of sap. . this water moves up because of atmospheric pressure during this period. water from the xylem vessel diffused into it. So that a sort of stair case movement of water took place in the plant.(B) Theory of Godlewski • He proposed that ascent of sap is brought about by periodic changes in osmotic pressure of xylem parenchyma and medullary rays. • Osmotic pressure of parenchyma increased. value of osmotic pressure of parenchyma decreased (lower down) and hence this water recieved by xylem which is lies at higher level. Thus it is known as "Relay pump theory" or "Clambering theory".

• He assumed that the pulsatory activity like heart beat is found in the inner cortical cells which are situated just above the endodermis.C. He proposed pulsation theory. water forced or pumped into cavities (Vessel) of xylem and water moves upward. Bose is known as father of Indian plant physiology.C. . Resulting of this pulsatory activity. • He conducted his experiment on a plant Desmodium gyrans (Indian telegraph plant) which is member of Papillionaceae family.(C) Pulsation theory of J. Bose • J.

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.• J. • According to him the pulsatory activity increased to 14 seconds by the application of narcotics drugs on the plant. .C.

. They proved by their experiment that the living cells are not essential for ascent of sap.Objections • The view of vital theory discarded by Straburger and Boucheries. Because when the living cells are killed (destroyed) by picric acid or HgCl2 solution even then ascent of sap continues.

(2) Root Pressure Theory • This theory was proposed by Priestley. . • Root pressure usually develops during the night when absorption is maximum and transpiration is minimum. • Root pressure : A positive pressure is present in the xylem sap of roots. • The term root pressure was coined by Stephan Hales. Priestley called it "hydrostatic pressure" Root pressure is also known as 'exudation pressure'. • The highest value of root pressure is found in those plants which are growing in well aerated and sufficient amount of soil during the humidity in the atmosphere.

This is also due to root pressure. • Guttation is also takes place due to root pressure. Therefore it is only applicable for small and herbaceous plants. • Ascent of sap through the root pressures only possible in small and herbaceous plants. .3 atmosphere. • The liquid which flows from the freshly decapitated stem of the plant is called bleeding.• The maximum value of root pressure falls in the range of 2 .

• 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. • (4) The rising up of water can not be possible at 2 atmosphere in tall plants. • (7) Root pressure takes place in the special conditions when the rate of absorption is higher and transpiration rate is minimum. . • (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. • Therefore. Upto 200 feet height plants require 10 atmosphere root pressure to push the water to the top of the plants. • (6) Root pressure is found less in summer and more in winter. theory explain partially the mechanism of ascent of sap. It is only effective in low transpiring herbaceous plants. root pressure is not significant in most of the plants. • Hence. • (5) A negative root pressure is found in some of the desert plants. • (3) The rate of ascent of sap is found rapidly even in the absence of root pressure.

(3) Physical Force Theory • According to this theory. the ascent of sap takes place by the vessels and tracheids through some physical forces. According to him the vessels and tracheid which are present in xylem. The following theories stated under this heading. • (A) Capillary force theory : • This theory was proposed by Boehm. . acts as capillaries and water rises up by surface tension in their capillaries.

But still ascent of sap occurs. . • Capillary theory is completely discarded on the basis of above reasons. • (b) The ends of vessels and tracheids are closed while ends of capillaries are open. But in tall plants the diameter of vessels and tracheids is more. tracheids are homologous as capillary. • (e) Lifting power of capillary is not much and can not account for the rise of water exceeding 3 meters.• Limitation of this theory as follows : • (a) Xylem vessels are not freely immersed into the water. • (d) Tall plants like Gymnosperms having tracheids instead of vessels which have many transverse septum. At this height capillary force does not operate. Therefore. It is capable for few centimeters rise only. • (c) The diameter of the capillaries should be very thin because capillary with narrow lumen encourages the entry of more water.

But now it has been clear that water rises up only through the lumen not by the walls. • Thus this theory also discarded. • The wall of vessels and tracheids of xylem are lignified and lignin is impermeable to water. • According to him water rises up in the wall of the xylem cells through the imbibition end through their lumen.(B) Imbibition force theory • This theory was proposed by Von Sachs. .

• Ascent of sap takes place due to spreading of air in the chain. • According to this theory. the molecules of water and air are arranged alternatively to form a chain. .(C) Chain theory • This theory proposed by Jamin.

cohesion force theory • This theory also known as "Water column theory" or "Cohesive force theory" • This theory was proposed by Dixon and Jolly.(D) Transpiration pull . • This is the most accepted theory at the present time and it accounts satisfactory explanation for the rising of water. This theory is based on the following facts : .

(A) Transpiration pull • Water is lost continuously from the leaves of the plants as a result of transpiration. Due to this mesophyll cells absorb water from adjoining internal mesophyll cells and compensate this loss of water. This pull is called "Negative pressure" because it is develops from aerial parts to the under ground part of the plants. Rapid transpiration develops a pull or tension in xylem which is called transpiration pull. Diffusion pressure deficit always remain higher in the mesophyll cells. As a result of loss of water from the mesophyll cell. Water vapour move out from the leaves. This loss ultimately compensating by xylem which leads water deficit in the xylem. . It is about 20 atmospheres.

.(b) Cohesion Force of Water • A force of mutual attraction present between the water molecules is called "Cohesive force". • Water molecules are held together continuously by cohesive force and to form a continuous water column. This cohesive force is up to 45 . 270 atmosphere.

• This water column pulled upwards continuously without breaking from the roots to the leaves by transpiration. • Both forces work together and maintain the continuity in between water and cell wall. .(c) Adhesive Force • Water molecules are also attached with the wall of vessels and tracheids through a force called adhesive force. • Cohesive force and adhesive force works continuously in the cavity (lumen) of xylem. Both the forces are responsible for maintaining unbroken continuity of water column from the roots to the leaves. • MacDougal called as "Hydrostatic system" of this continuous water column. Vessels and tracheids of xylem workds as pipe (Tubes).

• (3) The maximum value of osmotic pressure of the cells of leaves is found during noon. it absorb water inside the stem. • Maximum contraction is found during a maximum transpiration.• The process of ascent of sap a continues through the medium of above mechanisms. • (2) McDougal is found that daily contraction in stem of the trees with the help dendrograph apparatus. 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. • Evidences in support of Dixon's theory : • (1) Normally. Because the amount of water is minimum in that time . When this cut end is watered. This justifies the water tension in the stem.

Objection to Dixon's theory • (1) The presence of air bubbles in the xylem which may break the continuity of water column. • (2) Ascent of sap continues even after overlapping cuts. • * Removal of objections : .

Root pressure also helps to remove air bubbles. • (ii) Tracheid are also present along the vessels as alternative path. . but they are unable to move much distance because • (i) xylem vessels do not continue for long distances. pits are present between them. • (iii) Membrane and wall is present on the pits which is permeable. Pits are also present in them. Air bubbles may dissolve in water. but air bubbles do not pass out through them.• Although air bubbles are present in the xylem. to water.

• The origin of root pressure is an active process. • Factors affecting Ascent of sap • (1) Amount of water in soil. • Russel and Wooley found the ratio 50 : 1 by comparative study of water movement in apoplast and symplast.• The rate of transpiration is higher than that of rate of ascent of sap during the day time and is called absorption lag. (2) High temperature • (3) High atmospheric humidity (4) High atmospheric pressure • (5) Wind velocity (6) Number of stomata in leaves .

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Some proponents of vital theory are 1669-Ray-Sap Circulates in Plants like blood in Animals 1884-Godlewski-Cell pump water 1927-Bose-Sap ascends by pulsation of cortical movements 1938-Dixon and Barlee-Leaves Exude water by vital action of Leaf cells • • • • .Mechanism of Ascent of Sap • VITAL THEORY Suggestions have been made that living cells adjacent to conducting pipes of xylem are involved in the translocation of sap.These living cells are postulated to show some kind of pulsating action(just lioke heart beat) and pulsation of cells at successive intervals lifts the sap up.

Further numerous experiments have demonstrated clearly that living cells are not involved in the translocation of sap in the stem.• However anatomical studies have failed to show any pumping action in the Xylem. .

the positive pressure developed in the roots pumps the sap up in the xyulem ducts.For example.This theory seems to be applcable to many species.Grapevines have been shown to generate root pressure up to 5-6 bars.ROOT PRESSURE THEORY • Acc to Root pressure theory.which can support a water column upto 150-160 ft .which generate considerable amount of root pressure.

.it is low that it cannot account for upward translocation even upto a few ft. 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.• However.

.• 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.Because of Cohesive and adhesive forces.Water also shows forces of adhesive with the walls of xylem vessels and tracheids. .it exists as a column in the xylem elements.

Organismal Circulation Unicellular Organisms Autotrophic Multicellular Organisms (Heterotrophic Multicellular Organisms) .

mov The correct taxonomic affiliation is unclear. Slime molds can weigh up toward kilogram range and moves their blob-like mass around exclusively by cyclosis.botany. a slime mold This organism consists of one very large cytoplasm (plasmodium) with many nuclei and food vacuoles in the cytosol (coenocytic). Further study is needed to resolve its position. http://botit.Cyclosis in Physarum polycephalum. Transport is NOT always unidirectional. It has been treated as Fungus and Protist. What is the ATP source? . in a thin region of cytoplasm.edu/courses/img/Botany_1 30/Movies/Slime_mold.wisc. Here you can see. that it moves along pathways that are river-like in appearance.

Microfilaments of actin are found just under cell membrane.Cyclosis: cytoplasmic streaming…intracellular circulation Chloroplasts and other organelles have surface proteins with myosinlike activity. http://www.uk/mag/imgnov00/cycloa3i. Elodea canadensis ATP and Calcium allow myosin to slide along actin filaments. what would be the prediction on rate of cyclosis based on your hypothesis? .microscopy-uk.avi What is the source of ATP? Can you be more specific? If light intensity were reduced. QuickTime™ an d a Cinepak decompressor are need ed to see this p icture . resulting in circulation of organelles within the cell.org.

taking in CO2 and releasing O2 in daylight. But solutes need to be circulated in the large plant body as diffusion is too slow!! Node Internode Node Leaves Apical bud Axillary bud CO2 in and O2 out Branch O2 in and CO2 out Stem Lateral roots The root organ system is chemoheterotrophic. O2 in and CO2 out Taproot .Figure 36-3 Page 793 The shoot organ system is photoautotrophic. Diffusion is sufficient to exchange gases. taking in O2 and releasing CO2 in the darkness of the soil environment.

Branch Translocation Stem Transpiration The root system removes water and minerals from the soil environment. These solutes are transported to the roots in the phloem tissue: Node Internode Node Leaves Apical bud Axillary bud Carbohydrate etc.) by photosynthesis. These solutes are transported to the shoot in the xylem tissue: Translocation Lateral roots Transpiration Water and Minerals Taproot .Figure 36-3 Page 793 The shoot system produces carbohydrates (etc.

Because these pathways involve solutes in water passing in the adjacent tissues of a narrow vascular bundle. making a full circuit! Stem Transpiration Translocation Lateral roots Water and Minerals Taproot . and down the phloem.Figure 36-3 Page 793 Node Internode Apical bud Axillary bud Carbohydrate etc. this is a circulation system! Node Leaves Branch Transpiration and Translocation The water is moving up the xylem.

in part. rings of vascular bundles . concentric.Figure 36-18 Page 802 Plants occur in two major groups (and some minor ones) They differ. in their circulation systems: Cross section of a eudicot stem Cross section of a monocot stem Epidermis Cortex Pith Ground tissue Vascular bundles Dicots initially have one ring of vascular bundles Monocots rapidly develop multiple.

. new bundles are added toward the outside for increased circulation to the larger plant body.Monocot stem anatomy Mature Monocot Young Monocot vascular bundles As a monocot plant grows in diameter.

Norton & Company .Monocot stem anatomy Is this slice from a young or a mature part of the corn stem? Let’s take a closer look at the vascular tissues ©1996 Norton Presentation Maker. W. W.

W. Norton & Company Why must xylem do a lot more transport than phloem? .Monocot stem anatomy: vascular bundle Translocation Transpiration ©1996 Norton Presentation Maker. W.

W.Dicot circulation: stem anatomy Dicots start with one ring of bundles… Let’s take a closer look at the vascular tissues ©1996 Norton Presentation Maker. Norton & Company . W.

Dicot stem anatomy: vascular bundle phloem fibers Support of Stem functional phloem Translocation vascular cambium Cell Divison: More Xylem and Phloem xylem Transpiration As a dicot grows. W. Norton & Company . W. how does it add vascular capacity to become a tree? ©1996 Norton Presentation Maker.

Dicot stem anatomy: vascular cambium adds secondary tissues epidermis cortex 1º phloem 2º phloem cambium 2º xylem 1º xylem pith ©1996 Norton Presentation Maker. Norton & Company . W. W.

Norton & Company Each year the vascular cambium make a new layer of secondary xylem and secondary phloem .Dicot stem anatomy: vascular cambium adds secondary tissues ©1996 Norton Presentation Maker. W. W.

W.Dicot stem anatomy: four year-old stem (3 annual growth rings) phloem etc. Norton & Company . W. = bark All of these tissues were added by the vascular cambium! xylem = wood ©1996 Norton Presentation Maker.

Figure 36.29 Page 810 See also part (a) or less competition in forest? cambium phloem or more competition in forest? .

0 Page 791 periderm phloem cambium = bark heartwood pith .Figure 36.

The walls are strengthened with secondary thickenings including lignin. Norton & Company . tracheids suffice. W. W. Protoxylem have stretchable annular or helical thickenings. Metaxylem have reticulate or pitted and fully rigid walls. Tracheids have end walls and flow between cells is through pits. ©1996 Norton Presentation Maker. water needs are limited.Two Xylem Conducting Cells: tracheid developmental sequence Annular Helical Pitted When flowering plants are young.

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. 36. but lateral flow between cells is still through pits. Norton & Company .26 Page 806 As flowering plants age and grow. ©1996 Norton Presentation Maker. W.Two Xylem Conducting Cells: xylem vessel evolution plesiomorphic apomorphic Compare Fig. and tracheids need to be supplemented. W. water needs increase.

vessels. W. Norton & Company The huge vessel transports lots of water longitudinally. and tracheids ©1996 Norton Presentation Maker. and shows lots of pits for lateral transport . W.Dicot stem anatomy: xylem parenchyma.

W. and tracheids ©1996 Norton Presentation Maker.Dicot stem anatomy: xylem parenchyma. Norton & Company The huge vessel transports lots of water longitudinally. and shows lots of pits for lateral transport . vessels. W.

Norton & Company . W.Secondary xylem: cross sections of three different species Vessels. Tracheids have different distribution patterns. Some produce big vessels only in spring wood Others produce vessels year-round. W. ©1996 Norton Presentation Maker.

Norton & Company . O2 in and CO2 out ©1996 Norton Presentation Maker.Dicot stem anatomy: woody stem circulation 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. W. W.

. Rays transport water and minerals from the xylem to the exterior… …to keep the periderm. W. Norton & Company Rays transport sugar from the phloem toward the interior… …to keep pith and xylem parenchyma fueled.Dicot stem anatomy: 2-year old stem showing ray and periderm phloem ©1996 Norton Presentation Maker. and phloem parenchyma hydrated. cortex. W.

Xylem and Phloem: tissues with many cell types but conduction function toward pith main transpiration flow radial transport toward cortex main translocation flow ©1996 Norton Presentation Maker. W. W. Norton & Company .

nearctica. California 112 m tall (367.jpg .Mendocino Tree (Coastal Redwood) Sequoia sempervirens Ukiah.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.com/trees/conifer/tsuga/Ssemp10.

Transpiration in a tall tree has at least 3 critical components: Evaporation: pulling up water from above Capillarity: climbing up of water within xylem Root Pressure: pushing up water from below .

Transpiration: root pressure (osmotic “push”) Solutes from translocation of sugars accumulate in roots. Accumulating water in the root rises in the xylem. W. Water escapes from hydathodes. W. ©1996 Norton Presentation Maker. guttation This is not “dew” condensing! . Norton & Company Water from the soil moves in by osmosis.

fotocommunity. Root pressure accounts for maybe a half-meter of “push” up a tree trunk.Transpiration: root pressure (osmotic “push”) The veins (coarse and fine) show that no cell in a leaf is far from xylem and phloem (i.water and food!). http://img.e. The xylem of the veins leaks at the leaf margin in a modified stoma called the hydathode.jpg .com/photos/8489473. These droplets are xylem sap.

4m atmospheric pressure keeps water in tube Cohesion of water. 10.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”). helps avoid cavitation. caused by hydrogen bonds. A tree taller than 10.4 m would need some adaptations to avoid “cavitation” water .

W.Conifer stem anatomy: pine xylem tracheids with pits. xylem rays vascular cambium tracheids with pits In spite of the limitations of tracheidsonly xylem. W. conifers are among the tallest of trees! ray parenchyma ©1996 Norton Presentation Maker. Norton & Company .

As pressures change between adjacent cells. Norton & Company . the torus movement blocks catastrophic flow that would result in cavitation. ©1996 Norton Presentation Maker. W.Conifer stem anatomy: bordered pits as “check-valve” for flow P low P high These pit features allow conifers to be very tall and still avoid cavitation in their xylem cells. W.

Transpiration: evaporation (“pull”) 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 .

W.Grown in 32PO4 (radioactive phosphorus) 1 hour “Cold” medium 6 hours “Cold” medium 90 hours ©1996 Norton Presentation Maker. Norton & Company new growth black Is phosphate uptake from soil: transpiration or translocation? In xylem or phloem? Is phosphate mobilization from lower leaf: transpiration or translocation? In xylem or phloem? . W.

Translocation: How solutes move in phloem Leaf High Pressure plasmodesmata Root active transport Low Pressure Modified from: ©1996 Norton Presentation Maker. Norton & Company . W. W.

root tip Low Pressure Modified from: ©1996 Norton Presentation Maker. apical bud.Translocation: How solutes move bidirectionally in phloem Low Pressure Developing leaves. flowers fruits Leaf sugars amino acids High Pressure Lateral buds. Norton & Company . W. stems. roots. W.

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

Apical bud Axillary bud Carbohydrate etc.

Branch

Stem Transpiration

Translocation

Lateral roots

Water and Minerals

Taproot
Figure 36-3 Page 793

Node Internode Node Leaves

Apical bud Axillary bud Carbohydrate etc.

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

Water and Minerals

Taproot
Figure 36-3 Page 793

Thank you! Next Lecture: TRANSPIRATION .

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