OSMOSIS IN PLANT CELLS IS

INDIRECTLY ENERGY
DEPENDENT
Water, like any other substance, will only move down an energy gradient—that is, when there is a
difference in the energy of water in two parts of a system. In the case of the Figure 1.5, water
initially moves from chamber B to chamber A because the energy of pure water in chamber B is
greater than the energy of the water in the solution in chamber A. Net movement of water stops
when there is no longer an energy gradient across the selectively permeable membrane. Why is
the energy of pure water in chamber B of Figure 1.5 greater than the energy of the water in the
solution present in chamber A? The energy content of water, like any substance, is most easily
described in terms of its chemical potential.
Chemical potential () is defined as the free energy per mole of that substance and is a measure
of the capacity of a substance to react or move. The rule is that osmosis occurs only when there is
a difference in the chemical potential (_μ) of water on two sides of a selectively permeable membrane.
In other words, osmosis occurs only when the molar free energy of water, that is, the chemical
potential of water (w) on one side of a selectively permeable membrane, exceeds the molar free
energy or chemical potential of water on the other side of the same selectively permeable
membrane. It appears that the dissolution of a solute in water in some way affects the chemical
potential and hence the free energy of the solvent water molecules. Why is this so? This effect is
due to the fact that increasing solute concentration in an aqueous solution decreases the mole
fraction of water in the solution. The mole fraction of water (Xw) in a solution can be represented
as
Xw = w/w + s (1.2)
where w represents the moles of H2O in a given volume and s the moles of solute in the same
volume of solution. Thus, as one increases s at a constant value of w, the mole fraction of water,
Xw, decreases. Remember that the concentration of pure water is 55.5 moles L− 1. The chemical
potential of water (w) is related to the mole fraction of water (Xw) according to the following
equation
μw = μw
∗+ RT ln Xw (1.3)
where w* is the chemical potential of pure water under standard temperature and pressure, R is
the universal gas constant, and T is the absolute temperature. This equation tells us that as the
mole fraction of water decreases, the chemical potential and hence the molar free energy of water
decreases. Therefore, the higher the solute concentration of an aqueous solution, the lower the
chemical potential of the solvent water. Since energy flows ‘‘downhill’’ spontaneously, water flows
or diffuses spontaneously from chamber B to chamber A due to the difference in the chemical
potential of water between the two chambers separated by a selectively permeable membrane
(Figure 1.5).
Plant cells control the movement of water in and out of cells by altering the solute concentration
of the
cytosol relative to the solution external to the cell. One way to accomplish this is to regulate
transport of ions across the cell membrane into or out of the cell. For example, root cells can take
up nitrate ions NO− 3 from the soil by active transport to create a NO− 3 ion gradient across the
cell membrane (Figure 1.7) such that
(nitrate)in > (nitrate)out (1.4)
where μnitrate is the chemical potential of NO−
3 . Therefore,
it follows that
_μnitrate = (μnitrate)in − (μnitrate)out (1.5)
Since _μnitrate > 0, the uptake of NO−
3 is an active
transport process that requires an input of energy. Concomitantly,
the increase in cytosolic NO−
3 decreases the
mole fraction of water in the cytosol relative to the mole

THE CHEMICAL POTENTIAL OF WATER HAS AN OSMOTIC AS WELL AS A

PRESSURE COMPONENT

Osmosis can be easily demonstrated using a device known as an osmometer, constructed by

closing off the open end of a thistle tube with a selectively permeable membrane (Figure 1.8). If
the tube is then filled with a sugar solution and inverted in a volume of pure water, the volume of
solution in the tube will increase over time. The increase in volume is due to a net diffusion of
water across the membrane into the solution.
Osmosis dapat dengan mudah ditunjukkan dengan menggunakan alat yang dikenal sebagai osmometer, dibuat
dengan menutup ujung terbuka tabung thistle dengan membran permeabel yang selektif (Gambar 1.8). Jika
tabung kemudian diisi dengan larutan gula dan dibalikkan dengan volume air murni, volume larutan dalam
tabung akan meningkat seiring waktu. Peningkatan volume ini disebabkan oleh difusi bersih air melintasi
membran ke dalam larutan.

The increase in the volume of the solution will continue until the hydrostatic pressure developed
in the tube is sufficient to balance the force driving the water into the solution (Figure 1.8A).
Alternatively, the tube could be fitted with a piston that would allow us to measure the amount of
force required to just prevent any increase in the volume of solution (Figure 1.8B).
Peningkatan volume larutan akan berlanjut sampai tekanan hidrostatik yang dikembangkan dalam tabung cukup
untuk menyeimbangkan gaya yang mendorong air ke dalam larutan (Gambar 1.8A). Sebagai alternatif, tabung
dapat dilengkapi dengan piston yang memungkinkan kita untuk mengukur jumlah gaya yang diperlukan untuk
mencegah peningkatan volume larutan (Gambar 1.8B).

This force, measured in units of pressure (force per unit area), is known as osmotic pressure
(symbol = π; pi). The magnitude of the osmotic pressure that develops is a function of solute
concentration, at least in dilute solutions. It is useful to note that an isolated solution cannot have
an osmotic pressure. It has only the potential to manifest a pressure when placed in an
osmometer. For this reason, we say that the solution has an osmotic potential (symbol = _S). It is
convention to define osmotic potential as the negative of the osmotic pressure, since they are
equal but opposite forces(_S= − π).
Gaya ini, diukur dalam satuan tekanan (gaya per satuan luas), dikenal sebagai tekanan osmotik (simbol = π; pi).
Besarnya tekanan osmotik yang berkembang adalah fungsi konsentrasi zat terlarut, setidaknya dalam larutan
encer. Penting untuk dicatat bahwa solusi terisolasi tidak dapat memiliki tekanan osmotik. Ini hanya memiliki
potensi untuk mewujudkan tekanan ketika ditempatkan di osmometer. Untuk alasan ini, kami mengatakan
bahwa solusinya memiliki potensi osmotik (simbol = _S). Ini adalah konvensi untuk mendefinisikan potensi
osmotik sebagai negatif dari tekanan osmotik, karena mereka sama tetapi kekuatan berlawanan (_S = −π).

Whereas simple diffusion is driven entirely by differences in mole fractions of the solvent, it is
apparent from Figure 1.8 that pressure is also a factor in determining both the direction and rate
of water movement in an osmometer. Sufficient pressure applied to the piston (Figure 1.8B) will
prevent further net movement of water into the solution. If additional pressure were applied, we
might expect the net movement of water to reverse its direction and instead flow out of the
solution. Thus, osmosis is driven not only by the mole fraction of dissolved solute but by pressure
differences as well. Both of these factors influence the overall chemical potential of water, which is
the ultimate driving force for water movement in plants.
Sedangkan difusi sederhana sepenuhnya digerakkan oleh perbedaan fraksi mol pelarut, jelas dari Gambar 1.8
bahwa tekanan juga merupakan faktor dalam menentukan arah dan laju pergerakan air dalam osmometer.
Tekanan yang cukup diterapkan pada piston (Gambar 1.8B) akan mencegah pergerakan air lebih jauh ke dalam
larutan. Jika tekanan tambahan diterapkan, kita mungkin mengharapkan gerakan air bersih untuk membalikkan
arahnya dan sebaliknya mengalir keluar dari larutan. Jadi, osmosis tidak hanya didorong oleh fraksi mol dari zat
terlarut tetapi juga oleh perbedaan tekanan. Kedua faktor ini mempengaruhi potensi kimiawi air secara
keseluruhan, yang merupakan kekuatan pendorong utama pergerakan air pada tanaman.

The effect of pressure on chemical potential is represented by the value VP. V is the partial molal
volume, or the volume occupied by one mole of the chemical species. Since one liter (= 10− 3 m3)
of water contains 55.5 moles,Vw =(1000/55.5) or 18 ml per mole (= 18000mm3 mol− 1) of water. P is
the pressure. Measurements in plant physiology are commonly made at atmospheric pressure but
the presence of relatively rigid cell walls allows plant cells to develop significant hydrostatic
pressures. Hence it is convention to express P as the difference between actual pressure and
atmospheric pressure. The influence of pressure on chemicalpotential can now be added to
equation 1.3:
μw = μw∗ + RT ln Xw + VwP (1.7)
Pengaruh tekanan pada potensial kimia diwakili oleh nilai VP. V adalah volume molal parsial, atau volume yang
ditempati oleh satu mol spesies kimia. Karena satu liter (= 10−3 m3) air mengandung 55,5 mol, Vw = (1000 /
55,5) atau 18 ml per mol (= 18000mm3 mol − 1) air. P adalah tekanan. Pengukuran dalam fisiologi tanaman
biasanya dilakukan pada tekanan atmosfer tetapi keberadaan dinding sel yang relatif kaku memungkinkan sel-
sel tanaman untuk mengembangkan tekanan hidrostatik yang signifikan. Oleh karena itu, konvensi untuk
menyatakan P sebagai perbedaan antara tekanan aktual dan tekanan atmosfer. Pengaruh tekanan pada potensi
kimia sekarang dapat ditambahkan ke persamaan 1.3:

The chemical potential of water may also be influenced by electrical potential and gravitational
field. In spite of its strong dipole nature, the net electrical charge for water is zero and so the
electrical term can be ignored. This is why we talk about the chemical potential of water rather
than its electrochemical potential. While the gravitational term may be large and must be
considered where water movement in tall trees is concerned, it is not significant at the cellular
level.
Potensi kimiawi air juga dapat dipengaruhi oleh potensial listrik dan medan gravitasi. Terlepas dari sifat
dipolnya yang kuat, muatan listrik bersih untuk air adalah nol sehingga istilah listrik dapat diabaikan. Inilah
sebabnya kami berbicara tentang potensi kimiawi air daripada potensi elektrokimia. Sementara istilah gravitasi
mungkin besar dan harus dipertimbangkan dalam hal pergerakan air di pohon-pohon tinggi, itu tidak signifikan
pada tingkat sel.

Where water movement involves heights of 5 to 10 meters or less, the gravitational term is
commonly omitted. Since w = w* under very dilute conditions, it follows from Equation 1.7 that
PVw = − RT ln Xw (1.8)
Substituting π for P, we obtain the formal definition of osmotic pressure (Equation 1.10):
πVw = − RT ln Xw (1.9) or π = − RT ln XwVw − 1 (1.10)
As a matter of convenience, the value of Xw for pure water is arbitrarily designated as unity (= 1).
Consequently, Xw for a solution is always less than 1. Thus, equation 1.10 tells us that addition of
solute decreases the value of Xw, which in turn increases the osmotic pressure, π. By substituting
equation 1.9 in equation 1.7, we can now arrive at an expression relating the chemical potential of
water, hydrostatic pressure and osmotic pressure:
μw = μw ∗ + Vw(P − π) (1.11)

Jika pergerakan air melibatkan ketinggian 5 hingga 10 meter atau kurang, istilah gravitasi biasanya dihilangkan.
Karena w = w * dalam kondisi yang sangat encer, maka dari Persamaan 1.7 itu
PVw = − RT ln Xw (1.8)
Mengganti π untuk P, kita memperoleh definisi formal dari tekanan osmotik (Persamaan 1.10):
πVw = − RT ln Xw (1.9) or π = − RT ln XwVw − 1 (1.10)
Demi kenyamanan, nilai Xw untuk air murni secara sewenang-wenang ditetapkan sebagai satu kesatuan (= 1).
Akibatnya, Xw untuk solusi selalu kurang dari 1. Dengan demikian, persamaan 1.10 memberitahu kita bahwa
penambahan zat terlarut menurunkan nilai Xw, yang pada gilirannya meningkatkan tekanan osmotik, π. Dengan
mengganti persamaan 1.9 dalam persamaan 1.7, kita sekarang dapat sampai pada ekspresi yang berkaitan
dengan potensi kimiawi air, tekanan hidrostatik dan tekanan osmotik:
μw = μw ∗ + Vw(P − π) (1.11)

According to equation 1.11, the extent to which the chemical potential of water in a solution (μw)
differs
from that of pure water under conditions of standard temperature and pressure (μ∗w) is a function
of an osmotic component (π) and a pressure component (P).
Osmotic pressure or osmotic potential is one of the four colligative properties of a solution.
Since the magnitude of the osmotic pressure or osmotic potential exhibited by any solution is
dependent on the number of particles present in solution, osmotic pressure (π) and therefore
osmotic potential (_S) is a colligative property. Thus, a solution of 0.5M CaCl2 (Equation 1.12) will
exhibit a greater osmotic pressure than a solution of 0.5MNaCl (Equation 1.13) because the former
solution will exhibit twice as many Cl− ions present as the latter at the same concentration.

Menurut persamaan 1.11, sejauh mana potensi kimiawi air dalam larutan (μw) berbeda dari air murni dalam
kondisi suhu dan tekanan standar (μ ∗ w) adalah fungsi dari komponen osmotik (π) dan komponen tekanan (P).
Tekanan osmotik atau potensi osmotik adalah salah satu dari empat sifat koligatif suatu larutan. Karena
besarnya tekanan osmotik atau potensi osmotik yang diperlihatkan oleh larutan apa pun bergantung pada jumlah
partikel yang ada dalam larutan, tekanan osmotik (π) dan oleh karena itu potensi osmotik (_S) adalah sifat
koligatif. Dengan demikian, larutan 0,5M CaCl2 (Persamaan 1.12) akan menunjukkan tekanan osmotik yang
lebih besar daripada larutan 0,5MNaCl (Persamaan 1.13) karena solusi sebelumnya akan menunjukkan dua kali
lebih banyak ion Cl− yang hadir dengan yang terakhir pada konsentrasi yang sama.
CaCl2 → Ca2+ + 2Cl− (1.12)
NaCl → Na+ + Cl− (1.13)

The other three colligative properties of solutions are freezing point depression, boiling point
elevation, and the lowering of the vapor pressure of the solvent. Because π is directly proportional
to the solute molal concentration (moles of solute in 1 L of solvent), π can be used to calculate the
molecular mass of the solute molecule. According to the Ideal Gas Law,
PV = nRT (1.14)
where P is the pressure, V is the volume, n is the moles of the gaseous molecule, R is the
universal gas constant, and T is the absolute temperature. Thus, for an ideal, dilute solution
πV = nRT (1.15) or π = nRT V− 1 (1.16)
where π is the osmotic pressure of the solution, V is the volume, n is the moles of solute in the
solution, R is the universal gas constant, and T is the absolute temperature. Since n = mass
(m)/molecular mass (M), it follows that
π =mM− 1RT V− 1 (1.17)
Thus, by knowing the mass of the solute (m) dissolved in one liter (V− 1), one can measure π
experimentally for this solution and subsequently calculate the molecular mass of the solute.
Furthermore, if one knows the molecular mass of the solute, equation 1.17 allows one to calculate
the osmotic pressure (π) and hence the osmotic potential of a solution. Note that the solute must
be perfectly soluble in water in order to use this method to determine either the molecular mass
of the solute or the osmotic pressure of the solution.

Tiga sifat koligatif lainnya dari larutan adalah titik beku, peningkatan titik didih, dan penurunan tekanan uap
pelarut. Karena π berbanding lurus dengan konsentrasi molut terlarut (mol solut dalam 1 L pelarut), π dapat
digunakan untuk menghitung massa molekul dari molekul terlarut. Menurut Hukum Gas Ideal,
PV = nRT (1.14)
di mana P adalah tekanan, V adalah volumenya, n adalah mol molekul gas, R adalah konstanta gas universal,
dan T adalah suhu absolut. Ini, untuk solusi encer yang ideal
πV = nRT (1.15) or π = nRT V− 1 (1.16)
di mana π adalah tekanan osmotik larutan, V adalah volumenya, n adalah mol zat terlarut dalam larutan, R
adalah konstanta gas universal, dan T adalah suhu absolut. Karena n = massa (m) / massa molekul (M), maka ia
mengikuti itu
π =mM− 1RT V− 1 (1.17)
Dengan demikian, dengan mengetahui massa zat terlarut (m) yang dilarutkan dalam satu liter (V) 1), orang
dapat mengukur π secara eksperimental untuk larutan ini dan kemudian menghitung massa molekul zat terlarut.
Lebih lanjut, jika seseorang mengetahui massa molekul zat terlarut, persamaan 1.17 memungkinkan seseorang
untuk menghitung tekanan osmotik (π) dan karenanya potensi osmotik suatu larutan. Perhatikan bahwa zat
terlarut harus larut sempurna dalam air untuk menggunakan metode ini untuk menentukan massa molekul zat
terlarut atau tekanan osmotik dari larutan.