Phloem and Xylem

If you understand these two transport systems you have got the basics of fluid transport in plants. All vascular plants have both, though the cell form can vary (tracheids or vessel elements). Xylem is composed of dead, hollow cells (sclerenchyma) and is passive it carries the transpiration stream upwards from roots to leaves. Phloem cells are living and are used to transport sugars amino-acids and relocated minerals, typically from leaf to roots. In other words the opposite direction to xylem. Sugary solutions are actively pumped into the phloem from Source Cells and pumped out by Sink Cells. These are held side-by-side in linear complexes called vascular bundles.

The operation of Xylem

Xylem is dead hollow sclerenchyma cells which move water up the plant by transpiration tension cohesion. Transpiration: loss of water from the leaves (mainly through holes in the leaf surface called stomata. Tension the tensions that arise in the water columns of a leaf are transmitted down the water-bearing vessels of the whole tree. (This would fail if there were any large air bubbles, as the tension exerted may be many atmospheres). Dissolved gasses can come out of solution under this tension, and tree physiologists can put sensitive listening devices on tall trees in hot weather and hear tiny cracking noises as each microbubble appears). Cohesion the water column remains cohesive, transmitting the tension.

Pressures on xylem flow in a tree:

Evaporation. This imparts a suction pressure than puts immense tension onto the water columns in a truck trees shrink in diameter measurably on a hot sunny day. Guttation drops on leaf tips Roots push upwards. This can cause guttation (dripping of water from leaves resulting from active pumping), esp on damp mornings. The elevation attained is rarely more than a couple of metres. (Trees can drip sap in early spring).

Experiments on xylem flow

The German botanist Strasburger showed that 20m high trees if stood in tubs of poison (picric acid) would transport this to the tree top. Clearly this wasnt pumped!

Poison still sucked up to the top of the tree

Poison

sap

We can measure the tension in a twig using a pressure bomb: when sap is forced out of the twig the pressure in the sealed container = pressure in its xylem. Values are high enough to raise water 100m. High pressure

Symplastic vs Apoplastic flow

Big words, easy idea. This concerns the path taken by water when taken up from the soil. Does it go through cells cytoplasm, or in the spaces between cells? This turns out to matter: apoplastic means between cells, symplastic means going through cells plasma membranes. Root hair Root cortex Symplastic flow Apoplastic flow

This matters since the trans-membrane transport gives the plant the ability to filter and regulate the composition of its fluids.

The flow of water into roots is controlled by a band of corky, waterimpermeable cells lining the root cortex which force water to flow into the main vessels symplastically. This band of corky tissue (suberin + lignin) is the casparian strip, and is present in the endodermis of the root systems of most vascular plants. The casparian strip ensures that all water entering the stele of the root (thence up to the main stem) has passed through a plasma membrane so has been regulated by transport proteins. Casparian strip stele Cortex

Phloem
Phloem cells are involved in the active transport of sugars, amino acids and other metabolites around the plant. Their operation is very different to xylem: the main flow is downwards, and is inhibited by metabolic poisons. A simple if cruel experiment on this dates back to Malpighi around 1700. He girdled (ring-barked) trees and observed that the bark above the cut swelled while the bark below died (as eventually did the tree). This is because the cut stopped the downflow of metabolites in the outer regions of the bark (where trees phloem is found). We now know that phloem sap moves as fast as 1m per hour too fast for diffusion. Instead a form of bulk flow must be involved.

The model explaining how solutes are moved around the phloem is called the pressure flow model, and can be explained by consideration of osmosis, applied to solutions of two sugar solutions across a semipermeable membrane.

Water initially enters both ends by osmosis, but eventually the hydrostatic pressure on the semipermeable membrane offsets the osmotic pressure, stopping influx at the dilute (sink) end. The pressure is greater at the top end (where conc is higher), effectively pushing water into the conc end and out of the dilute end Water Initially influx at both ends Conc sugar (source end) Membrane bulges, imposing hydrostatic pressure Water Net flow of water and solutes along the tube

Dilute sugar (sink end) Water

Water