Water in plants

A. J. McElrone, Nature Education 2013

 What are the forces for ascent of water in plants?
Transpiration
Capillary forces Cohesion-tension

Microporous structure with diameter < 1µm Water-water molecule hydrogen bond
But flow rate very small
“reverse transpiration” during night, water uptake
from rain or dew through cracks in the leaf

Osmotic pressure

Transpiration-induced water loss, unbalanced

osmotic pressure

Interfacial forces

Vertically segmented xylem compartments

Bubbles adhered to the inner walls
Marangoni effect (high in halophytes)

Phil. Trans. R. Soc. Lond. B (1993)

 Osmotic contribution

Gibbs energy of solution Ideal solution

∆𝐺 = ∆𝐻 − 𝑇∆𝑆 ∆𝐻 = 0 𝑁!
Boltzmann
𝑆 = 𝑘ℓ𝑛𝑊 𝑁"
𝑁! + 𝑁" !
𝑆 = 𝑘ℓ𝑛
𝑁!! 𝑁"!
Sterling approximation
∆𝑆= - 𝑁!k ℓ𝑛𝑥!- 𝑁"k ℓ𝑛𝑥"

𝐺 = 0 𝑛# 𝜇#
#
𝜇 = 𝑐ℎ𝑒𝑚𝑖𝑐𝑎𝑙 𝑝𝑜𝑡𝑒𝑛𝑡𝑖𝑎𝑙
𝜇! = 𝜇!$+RT ℓ𝑛𝑥! 𝜇! = 𝜇!$+RT ℓ𝑛𝑥! 𝜇!$
Osmotic contribution to the ascent of water in plants (Transpiration)

Maximum height for water ascent

𝜇!""# + 𝑀𝑔ℎ!""# = 𝜇$%&' + 𝑀𝑔ℎ$%&' 𝑅𝑇 1

ℎ= ℓ𝑛
𝑀𝑔 𝑅𝐻
ℎ!""# = 0 RH = relative humidity
M = molecular weight
𝜇 ( + RTℓ𝑛𝑝( = 𝜇 ( + RTℓ𝑛(𝑅𝐻𝑝( ) + Mgh
leaf at 25oC and RH = 90%
root
1
ℎ = 1.40 10) ℓ𝑛
𝑅𝐻
𝑅𝐻𝑝( = actual partial pressure of water
vapor in the air
ℎ = 1475 𝑚
Assuming pool of pure water at the root

Science, 4265, 567, 1976

 Capillary-driven desalination

AAO/gel

Silica frit

Aquaporin Polyamide

A=Water permeability coefficient Soil Soil

2021
 https://www.youtube.com/watch?v=Q76jNr0z3s4

https://www.youtube.com/watch?v=Q76jNr0z3s4
Phrynosoma cornutum
“Liquid diode”: flow allowed in one direction but inhibited in the other
Capillary Rise
 Goodwin p. 206

Kinetics of capillary rise

𝚫P1 = Patm - Pliquid 1= Patm - Pliquid 0 - 𝝆gh

r = 𝚫Pyoung-laplace - 𝝆gh
R=
cosθ "8
𝚫Pyoung-laplace = cos𝜃
9

$&
∆𝑃0= %
cos𝜃-𝜌𝑔ℎ
Volumetric flow rate in a tube:

Hagen-Poiseuille Equation <9 "

#$
BCDEFGH> @>
𝑄= %
=>?
=𝜋𝑟 " @A
∆;! <9 " @>
𝑄= =𝜋𝑟 "
=>? @A
#$
9# BCDEFGH> @> !" $%&'()*+,-"%*
r = capillary radius
%
= !#
= ."/
=>? @A
h = wetted length of a capillary
 Goodwin p. 206

Kinetics of capillary rise

@> "98BCDEFGH>9 #
@A
= =>?
r
R=
cosθ @> 89BCDE GH9 #
@A
= I>? - =?

When the gravity term can be neglected

Washburn Equation FG IJKLMN h(t ) γ r cosθ t

FH
=
OGP ∫0
h dh =
4η ∫0
dt

J.K Application: Penetration of oils,

𝛾𝑟𝑐𝑜𝑠𝜃 water or surfactant relevant for
ℎ 𝑡 = 𝑡 drilling of oil and gas in recovery
4𝜂 from mature wells
 Capillary diode: Bioinspired inner microstructured tube controlled capillary rise

www.pnas.org/cgi/doi/10.1073/pnas.1821493116 (2019)
 Adamson, Cap. III

Thermodynamics of Liquid Interfaces

Effect of T on surface tension

Surface free energy per unit area

⎛ ∂G ⎞
GS = γ = ⎜
⎝ ∂A ⎟⎠ T , P
Thermodynamics of Liquid Interfaces Adamson, Cap. III

Close system during

phase separation

Clausius-Clapeyron
Pv
𝑑𝑃 𝐿
=
𝑑𝑇 𝑇∆𝑣
L = specific latent heat
Pl ∆𝑣 = specific
volume change of
phase transition Tc = critical temperature

2
𝑇
Guggenheim: 𝛾 = 𝛾1 1−
𝑇'

n ≃11/9 for organic liquids

Water
 Vapor pressure deficit (VPD):
Clausius-Clapeyron
Saturation vapor pressure - How much water is in the air
𝑑𝑃 𝐿
= 𝑃𝑉 = 𝑛𝑅𝑇 High T, high vapor pressure, high VPD
𝑑𝑇 𝑇∆𝑣
Water evaporates, drier soil
In atmospheric conditions
Plants draw more water from their roots, drier soil
@;&(()*) R, ;&(()*)
=
@Q SQ #

𝑃+(-&#) = saturation vapor pressure

/0.2*34
𝑑𝑃3()5#) = 6.1094𝑒 45*67.86

𝑑𝑃+(-&#) ≅ 6.1𝑒 (.(9:;<

Water holding capacity increases 7% for every 1oC

Journal of Applied Meteorology. 35 (4): 601–9 (1996)

 Yuan et al., Sci. Adv. 2019; 5: eaax1396

High VPD and plant death

VPD as drivers of the drought-related forest mortality.

High VPD may trigger stomatal closure to avoid excess water loss due to the high
evaporative demand of the air and tissue-level carbohydrate starvation. IPCC2021

Reduced soil water supply coupled with high evaporative demand causes xylem
conduits and the water channels to cavitate (become air-filled), stopping the flow of
water, desiccating plant tissues, and leading to plant death.
A warmer and more moist atmosphere over the oceans
makes it likely that the strongest hurricanes will be more
intense, produce more rainfall, and possibly be larger.

IPCC2021