1. Means of Transport — Overview

Plants transport substances across short distances (cell to cell) and long distances (root to shoot). Different mechanisms operate at each scale:

Mechanism Type Energy Required? Example
Diffusion Passive No O₂, CO₂ exchange in leaves; movement of ions across membranes
Facilitated Diffusion Passive (carrier/channel proteins) No Water through aquaporins; ions through ion channels
Osmosis Passive (special case of diffusion) No Water absorption by roots; stomatal movement
Active Transport Active (carrier proteins + ATP) Yes (ATP) Mineral ion uptake against concentration gradient

Apoplast and Symplast Pathways

Water moves through plant tissues via two routes:

  • Apoplast pathway: Movement through cell walls and intercellular spaces (outside the plasma membrane). Fast but blocked at the Casparian strip of endodermis.
  • Symplast pathway: Movement through the cytoplasm connected by plasmodesmata. Slower but continuous through living cells.
  • Transmembrane pathway: Water moves in and out of cells across membranes repeatedly — slowest pathway.

The Casparian strip (suberin band in endodermal cell walls) forces all apoplastic water to switch to the symplast pathway — this is where selective regulation of mineral uptake occurs.

2. Diffusion

Diffusion is the movement of molecules from a region of higher concentration to a region of lower concentration — down the concentration gradient — until equilibrium is reached.

  • Does not require a membrane — occurs in gases, liquids, and through membranes.
  • Rate depends on: concentration gradient, temperature, molecular size, and the medium.
  • Gases diffuse much faster than liquids.
  • In plants: CO₂ and O₂ exchange in leaves occurs by diffusion through stomata.

Facilitated Diffusion

Some molecules (e.g., water, charged ions) cannot cross the lipid bilayer directly — they use transport proteins:

  • Aquaporins: Channel proteins specific for water — allow rapid osmosis across membranes without energy expenditure.
  • Ion channels: Allow specific ions to move down their electrochemical gradient.
  • Carrier proteins: Bind specific molecules and change shape to transport them across.

Facilitated diffusion is still passive (no ATP) but is faster and more selective than simple diffusion.

3. Osmosis and Water Potential

Osmosis is the diffusion of water molecules across a selectively permeable membrane from a region of higher water potential to a region of lower water potential.

Water Potential (Ψ)

Water potential is the chemical potential of water — it determines the direction of water movement. Represented by the Greek letter psi (Ψ). Key points:

  • Pure water has the highest water potential = 0 (by convention).
  • Adding solutes lowers water potential (makes it more negative).
  • Water always moves from higher (less negative) to lower (more negative) water potential.

Ψ=Ψs+Ψp

where Ψs is the solute potential (osmotic potential) and Ψp is the pressure potential (turgor pressure).

Component Symbol Sign Effect on Water Potential
Solute Potential (Osmotic Potential) Ψs Always negative Lowers water potential; more solutes = more negative Ψs
Pressure Potential (Turgor Pressure) Ψp Usually positive (can be negative in xylem) Raises water potential; wall pressure increases Ψp
Water Potential Ψ Zero or negative (for solutions) Direction of water movement — high to low

Osmotic Terms — Turgid, Flaccid and Plasmolysed

Condition External Solution Cell State Ψp Cell Volume
Turgid Hypotonic (more dilute) Water enters; cell swells; wall exerts back pressure High (positive) Maximum
Flaccid Isotonic No net water movement; cell is limp Zero Normal
Incipient Plasmolysis Hypertonic (begin) Protoplast just begins to pull away from wall Zero Slightly reduced
Plasmolysed Hypertonic (concentrated) Protoplast shrinks away from cell wall; cell is plasmolysed Negative Reduced

Deplasmolysis

When a plasmolysed cell is placed in a hypotonic solution, water enters and the protoplast returns to its original position — this is called deplasmolysis. It is reversible if the cell has not been damaged. Plasmolysis is used to demonstrate selective permeability of the plasma membrane.

Osmotic Pressure vs Osmotic Potential

  • Osmotic pressure (OP): The pressure needed to prevent osmosis — always positive; higher concentration = higher OP.
  • Osmotic potential (Ψs): Always negative; numerically equal to OP but opposite in sign.
  • Ψs=OP

4. Imbibition

Imbibition is the absorption of water by solid hydrophilic colloids (called imbibants) causing them to swell. It is a special type of diffusion — the water moves along a water potential gradient but into a solid matrix.

  • Examples: dry seeds absorbing water and swelling; dry wood swelling when wet; agar plates absorbing water.
  • Imbibition generates enormous imbibitional pressure — sufficient to break hard seed coats and split rocks.
  • Requires an affinity between the imbibant and water (e.g., starch, proteins, cellulose are good imbibants).
  • Imbibition is the first step in seed germination — seeds imbibe water before metabolic activation begins.

5. Absorption of Water by Roots

The primary site of water absorption is the root hair zone — root hairs are extensions of epidermal cells with enormous surface area.

Mechanism of Water Absorption

  • Root hair cells have a lower water potential (due to dissolved solutes) than the soil water.
  • Water moves into root hairs by osmosis down the water potential gradient.
  • Water then moves inward from cell to cell (symplast or apoplast) towards the xylem.
  • The Casparian strip in the endodermis forces water into the symplast — allowing selective mineral uptake.

Active vs Passive Water Absorption

Type Driving Force Condition
Active absorption Root pressure (osmotic); metabolic energy used for mineral uptake which lowers water potential Slow transpiration; night; humid conditions
Passive absorption Transpiration pull — negative pressure (tension) in xylem pulls water up; no ATP used High transpiration; daytime; dominant mechanism

6. Transpiration

Transpiration is the loss of water vapour from aerial parts of plants (mainly through stomata). It is often called a "necessary evil" — wasteful in terms of water loss, but essential for creating the driving force for water movement.

Types of Transpiration

Type Site % of Total Transpiration
Stomatal transpiration Through stomata 80–90% (dominant)
Cuticular transpiration Through cuticle of leaves 5–10%
Lenticular transpiration Through lenticels of stem 1–5% (least)

Significance of Transpiration

  • Creates the transpiration pull — the main driving force for ascent of sap.
  • Helps in absorption and upward movement of water and minerals.
  • Cools the leaf surface by evaporative cooling.
  • Maintains the shape and structure of cells (turgor pressure).
  • Concentrates solutes in leaf cells, favouring photosynthesis.

Factors Affecting Transpiration

  • Temperature: Higher temperature increases transpiration (more evaporation).
  • Humidity: Higher humidity decreases transpiration (reduces the gradient).
  • Wind speed: Higher wind speed increases transpiration.
  • Light: Causes stomata to open → increases transpiration.
  • Water availability: Low water causes stomatal closure → reduces transpiration.
  • CO₂ concentration: High CO₂ causes stomatal closure → reduces transpiration.

Guttation

Guttation is the exudation of liquid water (not vapour) from hydathodes at leaf margins — typically in humid conditions at night/early morning when transpiration is low but root pressure is high. The liquid contains dissolved salts (unlike dew, which is pure condensed water).

7. Ascent of Sap — Movement of Water in Xylem

The upward movement of water and dissolved minerals through the xylem is called ascent of sap. In tall trees (up to 120 m), this requires tremendous force. Several theories have been proposed:

Theories of Ascent of Sap

Theory Proposed by Mechanism Status
Root Pressure Theory Priestley Osmotic pressure builds in roots pushing water up Insufficient alone — only pushes ~2–3 bar; cannot explain tall trees
Capillary Theory Surface tension in narrow xylem vessels pulls water up Insufficient — capillarity can only raise water ~1 m in xylem diameter
Cohesion-Tension-Transpiration Pull Theory Dixon & Joly (1894) Transpiration creates tension (negative pressure) in xylem; cohesion of water molecules transmits pull from leaves to roots Most widely accepted

Cohesion-Tension Theory — Details

Proposed by Dixon and Joly (1894) — also called the Transpiration Pull theory. It is based on three properties of water:

  • Transpiration pull: Water evaporates from mesophyll cells through stomata → water potential in mesophyll cells drops → water is drawn from xylem into mesophyll → negative pressure (tension) is created in xylem.
  • Cohesion of water: Water molecules are strongly attracted to each other by hydrogen bonds. This cohesion keeps the water column intact even under tension.
  • Adhesion: Water molecules adhere to the hydrophilic walls of xylem vessels — helps resist breaking of the water column.

The combination creates an unbroken water column from roots to leaves. The tension in xylem can reach −15 to −80 bar — sufficient to lift water in the tallest trees. This is passive transport — no metabolic energy is required.

Root Pressure

Root pressure is a positive pressure that develops in the root xylem due to active mineral uptake lowering the water potential of xylem. Water moves in by osmosis creating pressure. Evidence:

  • Bleeding: When a stem is cut, xylem fluid oozes out due to root pressure.
  • Guttation: Exudation of water droplets from hydathodes at night when transpiration is low.
  • Root pressure alone cannot account for tall trees — it supplements the transpiration pull.

8. Opening and Closing of Stomata

Stomata are pores in the leaf epidermis surrounded by two kidney-shaped (or dumbbell-shaped in grasses) guard cells. Their opening and closing regulates gas exchange and transpiration.

Mechanism of Stomatal Opening

  • In light, guard cells photosynthesise — CO₂ is used up, pH rises.
  • Starch is converted to malate and glucose — osmotic concentration increases.
  • K⁺ ions are actively pumped into guard cells — water potential decreases.
  • Water enters guard cells by osmosis → guard cells become turgid → stomata open.
  • The inner wall of guard cells is thicker and inextensible — so turgidity causes the cells to bow outward, opening the pore.

Mechanism of Stomatal Closing

  • In darkness or water stress: K⁺ ions move out of guard cells.
  • Osmotic concentration decreases → water moves out by osmosis.
  • Guard cells become flaccid → stomata close.
  • ABA (Abscisic Acid) triggers stomatal closure under water stress — acts as the stress signal.
Condition Stomatal Response Mechanism
Light Open K⁺ influx, malate accumulation, guard cells turgid
Darkness Close K⁺ efflux, guard cells flaccid
Water deficit (drought) Close ABA release triggers K⁺ efflux
High CO₂ Close CO₂ accumulation inhibits K⁺ uptake
Low CO₂ Open Promotes K⁺ influx into guard cells