Transpiration and Translocation

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Biology:

life study of

What is Life?

Properties of Life

Cellular Structure: the unit of life, one or many

Metabolism: photosynthesis, respiration, fermentation, digestion, gas exchange, secretion, excretion, circulation--processing materials and energy

Growth: cell enlargement, cell number

Movement: intracellular, movement, locomotion

Reproduction: avoid extinction at death

Behavior: short term response to stimuli

Evolution: long term adaptation

Organismal Circulation

Unicellular Organisms

Autotrophic Multicellular Organisms

(Heterotrophic Multicellular Organisms)

Cyclosis in Physarum polycephalum , a slime mold

This organism consists of one very large cytoplasm

(plasmodium) with many nuclei and food vacuoles in the cytosol

(coenocytic).

Slime molds can weigh up toward kilogram range and move their blob-like mass around exclusively by cyclosis.

http://botit.botany.wisc.edu/courses/img/Botany_

130/Movies/Slime_mold.mov

Here you can see, in a thin region of cytoplasm, that it moves along pathways that are river-like in appearance.

Transport is NOT always unidirectional.

The correct taxonomic affiliation is unclear.

It has been treated as

Fungus and Protist.

Further study is needed to resolve its position.

What is the ATP source?

Cyclosis: cytoplasmic streaming…intracellular circulation

Elodea canadensis

Chloroplasts and other organelles have surface proteins with myosin-like activity.

Microfilaments of actin are found just under cell membrane.

http://www.microscopy-uk.org.uk/mag/imgnov00/cycloa3i.avi

What is the source of ATP?

ATP and

Calcium allow myosin to slide along actin filaments, resulting in circulation of organelles within the cell.

Can you be more specific?

If light intensity were reduced, what would be the prediction based on your hypothesis?

Figure 36-3 Page 793

The shoot organ system is photoautotrophic, taking in CO

2 releasing O daylight.

2 and in

Diffusion is sufficient to exchange gases.

But solutes need to be circulated in the large plant body as diffusion is too slow!!

The root organ system is chemoheterotrophic, taking in O

2 and releasing CO

2 in the darkness of the soil environment.

Node

Internode

Node

Leaves

Stem

Apical bud

Axillary bud

CO

2 in and O

2 out

Branch

O

2 in and CO

2 out

Lateral roots

Taproot

O

2 in and CO

2 out

Figure 36-3 Page 793

The shoot system produces carbohydrates (etc.) by photosynthesis.

These solutes are transported to the roots in the phloem tissue:

Translocation

Node

Internode

Node

Leaves

Apical bud

Axillary bud

Carbohydrate etc.

Branch

Stem

Transpiration Translocation

Lateral roots

The root system removes water and minerals from the soil environment. These solutes are transported to the shoot in the xylem tissue:

Transpiration

Water and

Minerals

Taproot

Figure 36-3 Page 793

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!

Node

Internode

Node

Leaves

Apical bud

Axillary bud

Carbohydrate etc.

Branch

Stem

Transpiration Translocation

Lateral roots

Water and

Minerals

Taproot

Figure 36-18 Page 802

Plants occur in two major groups (and some minor ones)

They differ, in part, 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, concentric, rings of vascular bundles

Monocot circulation: transpiration and translocation

©1996 Norton Presentation Maker, W. W. Norton & Company

Monocot stem anatomy

Young Monocot

Mature Monocot vascular bundles

As a monocot plant grows in diameter, new bundles are added toward the outside for increased circulation to the larger plant body.

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. Norton & Company

Monocot stem anatomy: vascular bundle

Translocation

Transpiration

©1996 Norton Presentation Maker, W. W. Norton & Company

Why must xylem do a lot more transport than phloem?

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, W. W. Norton & Company

Dicot stem anatomy: vascular bundle

©1996 Norton Presentation Maker, W. W. Norton & Company phloem fibers

Support of Stem functional phloem

Translocation vascular cambium

Cell Divison: More

Xylem and Phloem xylem Transpiration

As a dicot grows, how does it add vascular capacity to become a tree?

Dicot stem anatomy: vascular cambium adds secondary tissues epidermis cortex

1º phloem

2º phloem cambium

2º xylem

1º xylem pith

Dicot stem anatomy: vascular cambium adds secondary tissues

©1996 Norton Presentation Maker, W. W. Norton & Company

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

©1996 Norton Presentation Maker, W. W. Norton & Company

Figure 36.29 Page 810

See also part (a) cambium phloem or less competition in forest?

or more competition in forest?

Figure 36.0 Page 791 periderm phloem cambium

= bark heartwood pith

Dicot stem anatomy: 2-year old stem showing ray and periderm

©1996 Norton Presentation Maker, W. W. Norton & Company

Dicot stem anatomy: periderm dying epidermis maturing cork cells periderm cork cambium phelloderm cortical collenchyma cortical parenchyma

©1996 Norton Presentation Maker, W. W. Norton & Company

Two Xylem Conducting Cells: tracheid developmental sequence

Annular Helical Pitted

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.

Metaxylem have reticulate or pitted and fully rigid walls.

Tracheids have end walls and flow between cells is through pits.

©1996 Norton Presentation Maker, W. W. Norton & Company

Dicot stem anatomy: xylem vessel evolution plesiomorphic apomorphic

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.

©1996 Norton Presentation Maker, W. W. Norton & Company

Dicot stem anatomy: xylem parenchyma, vessels, and tracheids

©1996 Norton Presentation Maker, W. W. Norton & Company

Dicot stem anatomy: xylem parenchyma, vessels, and tracheids

The huge vessel transports lots of water longitudinally, and shows lots of pits for lateral transport

Dicot stem anatomy: woody stem circulation

O

2 in and CO

2 out

©1996 Norton Presentation Maker, W. W. Norton & Company

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.

Secondary xylem: cross sections of three species

Vessels, Tracheids have different distribution patterns.

Some produce big vessels only in spring wood

Others produce vessels year-round.

Xylem and Phloem: tissues with many cell types but conduction function

©1996 Norton Presentation Maker, W. W. Norton & Company

Mendocino Tree (Coastal Redwood) Sequoia sempervirens

Ukiah, California

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 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 ” ) guttation

This is not “ dew ” condensing!

Solutes from translocation of sugars accumulate in roots.

Water from the soil moves in by osmosis.

Accumulating water in the root rises in the xylem.

Water escapes from hydathodes.

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.

http://img.fotocommunity.com/photos/8489473.jpg

These droplets are xylem sap.

Root pressure accounts for maybe a half-meter of “ push ” up a tree trunk.

Capillarity: maximum height of unbroken water column gravity pulls water down atmospheric pressure keeps water in tube glass tube vacuum created

10.4m

The small diameter of vessels and tracheids and the surface tension of water provide capillary

( “ climb ” ).

Cohesion of water, caused by hydrogen bonds, helps avoid cavitation.

water

A tree taller than

10.4 m would need some adaptations to avoid “ cavitation ”

Dicot stem anatomy: pine xylem tracheids with pits, xylem rays tracheids with pits ray parenchyma

©1996 Norton Presentation Maker, W. W. Norton & Company

In spite of the limitations of tracheids-only xylem, conifers are among the tallest of trees!

Conifer stem anatomy: bordered pits as “ check-valve ” for flow secondary wall primary wall middle lamella pit aperture pit membrane pit border torus pit chamber

These pit features allow conifers to be very tall and still avoid cavitation in their xylem cells.

P low

P high

As pressures change between adjacent cells, the torus movement blocks catastrophic flow that would result in cavitation.

Transpiration: evaporation ( “ pull ” )

Water evaporating from a porous clay cap also lifts the mercury!

water mercury

Transpiration can lift the mercury vacuum above its normal cavitation height!

76 cm mercury

Grown in 32 PO

4

(radioactive phosphorus) 1 hour

“ Cold ” medium 6 hours “ Cold ” medium 90 hours 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?

Translocation: How solutes move in phloem

Leaf High Pressure plasmodesmata

Root

Low Pressure

Modified from: ©1996 Norton Presentation Maker, W. W. Norton & Company

Translocation: How solutes move bidirectionally in phloem

Low Pressure

Developing leaves, apical bud, flowers fruits

Leaf sugars amino acids

High

Pressure

Low Pressure

Modified from: ©1996 Norton Presentation Maker, W. W. Norton & Company

Lateral buds, stems, roots, root tip

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

Water and

Minerals

Apical bud

Axillary bud

Taproot

Figure 36-3 Page 793

Carbohydrate etc.

Branch

Stem

Transpiration Translocation

Lateral roots

Node

Internode

Node

Leaves

Stem

Transpiration

Water and

Minerals

Apical bud

Axillary bud

Carbohydrate etc.

Branch

Translocation

Taproot

Figure 36-3 Page 793

Lateral roots

Translocation

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).

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