Aristotle (384 BC - 322 BC) is credited with the traditional classification of the five sense

organs: sight, smell, taste, touch, and hearing. As far back as the 1760's, the famous
philosopher Immanuel Kant proposed that our knowledge of the outside world depends on
our modes of perception. In order to define what is "extrasensory" we need to define what
is "sensory". Each of the 5 senses consists of organs with specialized cellular structures that
have receptors for specific stimuli. These cells have links to the nervous system and thus to
the brain. Sensing is done at primitive levels in the cells and integrated into sensations in
the nervous system. Sight is probably the most developed sense in humans, followed
closely by hearing.

Sight.

The eye is the organ of vision. It has a complex structure consisting of a transparent lens
that focuses light on the retina. The retina is covered with two basic types of light-sensitive
cells-rods and cones. The cone cells are sensitive to color and are located in the part of the
retina called the fovea, where the light is focused by the lens. The rod cells are not sensitive
to color, but have greater sensitivity to light than the cone cells. These cells are located
around the fovea and are responsible for peripheral vision and night vision. The eye is
connected to the brain through the optic nerve. The point of this connection is called the
"blind spot" because it is insensitive to light. Experiments have shown that the back of the
brain maps the visual input from the eyes.

The brain combines the input of our two eyes into a single three-dimensional image. In
addition, even though the image on the retina is upside-down because of the focusing action
of the lens, the brain compensates and provides the right-side-up perception. Experiments
have been done with subjects fitted with prisms that invert the images. The subjects go
through an initial period of great confusion, but subsequently they perceive the images as
right side up.

The range of perception of the eye is phenomenal. In the dark, a substance produced by the
rod cells increases the sensitivity of the eye so that it is possible to detect very dim light. In
strong light, the iris contracts reducing the size of the aperture that admits light into the eye
and a protective obscure substance reduces the exposure of the light-sensitive cells. The
spectrum of light to which the eye is sensitive varies from the red to the violet. Lower
electromagnetic frequencies in the infrared are sensed as heat, but cannot be seen. Higher
frequencies in the ultraviolet and beyond cannot be seen either, but can be sensed as
tingling of the skin or eyes depending on the frequency. The human eye is not sensitive to
the polarization of light, i.e., light that oscillates on a specific plane. Bees, on the other
hand, are sensitive to polarized light, and have a visual range that extends into the
ultraviolet. Some kinds of snakes have special infrared sensors that enable them to hunt in
absolute darkness using only the heat emitted by their prey. Birds have a higher density of
light-sensing cells than humans do in their retinas, and therefore, higher visual acuity.

Color blindness or "Daltonism" is a common abnormality in human vision that makes it

impossible to differentiate colors accurately. One type of color blindness results in the
inability to distinguish red from green. This can be a real handicap for certain types of
occupations. To a colorblind person, a person with normal color vision would appear to have
extrasensory perception. However, we want to reserve the term "extrasensory perception"
for perception that is beyond the range of the normal.

Click here for a Color Blindness test

and some amazing optical illusions. 

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Hearing.

The ear is the organ of hearing. The outer ear protrudes away from the head and is shaped
like a cup to direct sounds toward the tympanic membrane, which transmits vibrations to
the inner ear through a series of small bones in the middle ear called
the malleus, incus and stapes. The inner ear, or cochlea, is a spiral-shaped chamber
covered internally by nerve fibers that react to the vibrations and transmit impulses to the
brain via the auditory nerve. The brain combines the input of our two ears to determine the
direction and distance of sounds.
The inner ear has a vestibular system formed by three semicircular canals that are
approximately at right angles to each other and which are responsible for the sense of
balance and spatial orientation. The inner ear has chambers filled with a viscous fluid and
small particles (otoliths) containing calcium carbonate. The movement of these particles
over small hair cells in the inner ear sends signals to the brain that are interpreted as
motion and acceleration.

The human ear can perceive frequencies from 16 cycles per second, which is a very deep
bass, to 28,000 cycles per second, which is a very high pitch. Bats and dolphins can detect
frequencies higher than 100,000 cycles per second. The human ear can detect pitch
changes as small as 3 hundredths of one percent of the original frequency in some
frequency ranges. Some people have "perfect pitch", which is the ability to map a tone
precisely on the musical scale without reference to an external standard. It is estimated that
less than one in ten thousand people have perfect pitch, but speakers of tonal languages
like Vietnamese and Mandarin show remarkably precise absolute pitch in reading out lists of
words because pitch is an essential feature in conveying the meaning of words in tone
languages. The Eguchi Method teaches perfect pitch to children starting before they are 4
years old. After age 7, the ability to recognize notes does not improve much.

Taste.

The receptors for taste, called taste buds, are situated chiefly in the tongue, but they are
also located in the roof of the mouth and near the pharynx. They are able to detect four
basic tastes: salty, sweet, bitter, and sour. The tongue also can detect a sensation called
"umami" from taste receptors sensitive to amino acids. Generally, the taste buds close to
the tip of the tongue are sensitive to sweet tastes, whereas those in the back of the tongue
are sensitive to bitter tastes. The taste buds on top and on the side of the tongue are
sensitive to salty and sour tastes. At the base of each taste bud there is a nerve that sends
the sensations to the brain. The sense of taste functions in coordination with the sense of
smell. The number of taste buds varies substantially from individual to individual, but
greater numbers increase sensitivity. Women, in general, have a greater number of taste
buds than men. As in the case of color blindness, some people are insensitive to some
tastes.
Smell.

The nose is the organ responsible for the sense of smell. The cavity of the nose is lined with
mucous membranes that have smell receptors connected to the olfactory nerve. The smells
themselves consist of vapors of various substances. The smell receptors interact with the
molecules of these vapors and transmit the sensations to the brain. The nose also has a
structure called the vomeronasal organ whose function has not been determined, but which
is suspected of being sensitive to pheromones that influence the reproductive cycle. The
smell receptors are sensitive to seven types of sensations that can be characterized as
camphor, musk, flower, mint, ether, acrid, or putrid. The sense of smell is sometimes
temporarily lost when a person has a cold. Dogs have a sense of smell that is many times
more sensitive than man's.

Touch.

The sense of touch is distributed throughout the body. Nerve endings in the skin and other
parts of the body transmit sensations to the brain. Some parts of the body have a larger
number of nerve endings and, therefore, are more sensitive. Four kinds of touch sensations
can be identified: cold, heat, contact, and pain. Hairs on the skin magnify the sensitivity and
act as an early warning system for the body. The fingertips and the sexual organs have the
greatest concentration of nerve endings. The sexual organs have "erogenous zones" that
when stimulated start a series of endocrine reactions and motor responses resulting in
orgasm.

Beyond the five sense organs.

In addition to sight, smell, taste, touch, and hearing, humans also have awareness of
balance (equilibrioception), pressure, temperature (thermoception), pain (nociception), and
motion all of which may involve the coordinated use of multiple sensory organs. The sense
of balance is maintained by a complex interaction of visual inputs, the proprioceptive
sensors (which are affected by gravity and stretch sensors found in muscles, skin, and
joints), the inner ear vestibular system, and the central nervous system. Disturbances
occurring in any part of the balance system, or even within the brain's integration of inputs,
can cause the feeling of dizziness or unsteadiness.

Kinesthesia is the precise awareness of muscle and joint movement that allows us to
coordinate our muscles when we walk, talk, and use our hands. It is the sense of
kinesthesia that enables us to touch the tip of our nose with our eyes closed or to know
which part of the body we should scratch when we itch.

Synesthesia.

Some people experience a phenomenon called synesthesia in which one type of stimulation
evokes the sensation of another. For example, the hearing of a sound may result in the
sensation of the visualization of a color, or a shape may be sensed as a smell. Synesthesia
is hereditary and it is estimated that it occurs in 1 out of 1000 individuals with variations of
type and intensity. The most common forms of synesthesia link numbers or letters with
colors.

Breathing.

The very act of breathing, which we have to do several times per minute, brings air into our
lungs and whatever else is in the air. Our nose hairs filter out insects and some dust, but
gases like carbon monoxide, vapors, smoke, pollens, bacteria, viruses, and small dust
particles are carried into the lungs. The body has many self-cleansing mechanisms to keep
the lungs clean, but constant exposure to air pollutants eventually take their toll on the
lungs or other organs of the body. The tar and chemicals carried in the smoke of cigarettes
has been linked to many types of respiratory disorders and nicotine has an addictive effect
on the brain. Carbon monoxide produced by gasoline motors or charcoal fires in enclosed
places interferes with the oxygen-carrying function of the blood and is responsible for many
deaths each year. Paint solvents and gasoline fumes can damage the liver. Gases like
nitrous oxide and vapors like ether affect the nervous system and are used as anesthetics.

Eating and drinking.

We have to eat and drink to sustain our life, but what we ingest can carry not only
nutrients, but also substances that can adversely affect our health and mental processes.
There are regions in the United States where there is a great prevalence of kidney stones
that are associated with the hardness of the water. The "goiter belt" is another region where
the soil has a deficiency of iodine that would result in thyroid gland problems were it not for
iodized salt. Grain tainted with ergot fungus, which has an LSD component, has been
theorized to have caused hallucinations responsible for the witch hunts in Salem,
Massachusetts.

We cook foods to make them more digestible and to kill harmful microorganisms and
parasites. However, cooking may decrease the nutritional value of the food and charring
during grilling may create nitrosamines that have been associated with some cancers. Our
mass markets require the preservation of food by the use of food additives. Many of these
preservatives were discovered by analyzing foods, such as cheese, which don't readily spoil.
Other food additives are only used to improve appearance, e.g., artificial colors. Not all
additives are harmful, but some people prefer to buy "natural" or "organic" products
because they do not want to eat residual pesticides used in agriculture. Some "natural" and
"organic" products may be quite harmful. Opium, coca leaves, marijuana, and tobacco are
all natural products with addictive or mind-altering properties. This does not mean that they
may not have legitimate medical uses. Opium has been the source of morphine, which is a
powerful pain killer.

Caffeine, which occurs naturally in coffee, tea, and chocolate, is a nervous system stimulant
and also a diuretic. Large amounts of caffeine can cause tremors or shaking. Caffeine can
be addictive for some people even in small amounts. It is not by coincidence that soft drink
manufacturers use caffeine as an additive. If you drink more than one cup of coffee, tea,
chocolate, or cola drink per day you may be addicted to caffeine. This can be easily verified
by abstaining from caffeine-containing foods or drinks for a couple of days. Restlessness,
sinus pressure, or headaches are common withdrawal symptoms.

Medicines and drugs.

Medicines and drugs may be administered orally, by injection, inhalation, etc. The purpose
of medicines is to help the organism return to a healthy state. However, sometimes
medicines are prescribed to maintain a "normal" state. Antibiotics fall into the first category.
Once an infection has been eliminated, the medication can be stopped. Diabetes is in the
second category. It is necessary to take insulin all your life in order to live normally. With
the large number of drugs available, it is not surprising to find that some of them interact or
interfere with each other. Some women on birth control pills have become pregnant while
taking some types of antibiotics. Also, grapefruit has been found to elevate levels of some
medicines to toxic levels.

"Recreational" or illegal drugs such as heroin, cocaine, and LSD are mind-altering drugs that
affect the brain adversely, sometimes permanently. Certain non-prescription medicines,
such as cough suppressant syrups with dextromethorphan, act on the brain and can dull
thinking and creative abilities. "Ritualistic drugs", such as peyote, have hallucinogenic
properties and are used in certain religious ceremonies. Alcohol is the most frequently
abused mind-dulling drug. It acts as a brain intoxicant that reduces reaction times and
impairs the motor functions of the body. Drugs used in psychiatry also modify the way in
which the brain works. When used to treat depression or other debilitating mental
conditions these drugs actually help to restore the normal functions of the brain, but
generally not without side effects.

Skin absorption.
The skin acts as a protective barrier for the body, but it is not impervious. Many substances
pass through the skin and can affect various organs of the body. When the skin is exposed
to harsh chemicals, such as chlorine bleach or detergents, there may be just a local
irritation or chemical burn. Organic solvents such as gasoline, mineral spirits, and dry
cleaning fluids can be absorbed through the skin and reach toxic levels in the body. The
liver is the organ most frequently damaged as it tries to detoxify these substances.

Radiation/Light.

Electromagnetic radiation can be good and it can be bad for the body. It depends on the
type of the radiation and the duration of the exposure. Infrared radiation, which is low-
frequency radiation, is felt as heat. Sitting by a fireplace or a pot-bellied stove on a cold
winter night can feel comforting without any harmful effects. Excessive doses of infrared
radiation can result in burns. Normal skin produces Vitamin D when exposed to sunlight for
brief periods of time. When exposed for long periods of time, the skin reddens and becomes
painful to the touch. Repeated exposure to sunlight stimulates the skin to produce a
protective dark pigment called melanin. Chronic exposure to sunlight eventually breaks
down the cellular structure of the skin and can result in wrinkling, cancerous melanomas, or
other skin disorders.

The amount of light to which the body and eyes are exposed may affect the central nervous
system. Seasonal Affective Disorder (SAD) is a form of depression that includes feelings of
sadness, tiredness and cravings for carbohydrates. It is believed to be related to the
decreased sunlight in winter and the release of melatonin. Melatonin is usually produced by
the pineal gland at night and induces sleep. Besides sunlight, the body may also be exposed
to moonlight, which is sunlight reflected off the moon. The light of the moon is weak, but it
enabled early humans to have some nighttime activities before the invention of fire and
artificial lighting. The word "lunatic" is derived from the Latin for "moon". At one time it was
believed that the influence of the moon triggered mental disorders.

The use of artificial lighting has introduced some problems that did not exist before its
invention. Some people are sensitive and can get headaches from the flickering of
fluorescent lights. The flickering is particularly noticeable in the peripheral vision. Flashing
images from television or strobe lights can also cause harmful effects to the nervous system
and can trigger seizures. A Japanese television cartoon program that used flashing pictures
to simulate an explosion sent several hundred children to the hospital with various
neurological symptoms.

High-energy radiation such as ultraviolet light, X-rays, or Gamma rays can destroy cells. X-
rays and Gamma rays have greater penetration than ultraviolet light and are used medically
for diagnostic imaging and to burn tumors. Ultraviolet lights, also called "black" lights, are
used in hospitals and grocery stores to kill bacteria, but sometimes they are misused for
entertainment in bars or other dark places because ultraviolet light makes some substances
fluoresce.

Cosmic rays, which are high-energy particles, normally do not penetrate the earth's
atmosphere. However, astronauts have reported seeing flashes of light that have been
attributed to the effects of cosmic rays either on the eyes or the visual cortex of the brain.

Sounds.
Our bodies respond to sounds in fairly mechanical ways. Sudden noises can cause a person
to jump away from the noise, or turn the head in the direction of the noise. Soothing,
rhythmic noises such as the sound of the sea, a gurgling brook, or the beating heart in a
mother's breast are well known for their calming effects. Buzzing sounds close to the ears
cause us to wave the hands by our ears as if to repel insects. Certain high-pitched noises
such as scratching fingernails on a blackboard or the noise of a pencil on paper can "make
your skin crawl", which is an erection of the hairs on the skin. Loud repeated noises can
reduce the sensitivity of the ears and eventually cause hardness of hearing or even
deafness. Boilermakers that used noisy riveting equipment were particularly prone to
deafness as an occupational hazard. In the era of high-fidelity sound equipment with
powerful amplifiers, many young people are losing their hearing by listening to music at
very loud levels.

Bacteria and Viruses.

Bacteria and viruses come into the body principally through the eyes, the mouth, the skin,
and the nose. Some bacteria actually have a beneficial effect. The "normal flora" that are
found in the mouth release substances that prevent more harmful bacteria from getting
established. Other bacteria aid in digestion or produce vitamins and nutrients that the body
can use. Bread, yogurt, beer, wine, vinegar, and many types of cheeses are produced by
using specific types of non-harmful yeasts, bacteria, or fungi.

Disease-causing bacteria release toxins that interfere with normal body processes. Viruses,
which are much smaller than bacteria, work against the body by re-directing the synthesis
of normal cell components into replication of the virus. The body tries to fend off bacteria
and viruses by generating chemical antibodies and by increasing body temperature. Fever
creates a more hostile environment for bacteria but can result in delirium and other forms of
mental changes. Some diseases like rabies or polio attack directly the nervous
system. Learn more about Bacteria and Viruses.

Insect bites and stings.

Insect bites and stings are unpleasant inputs to the human body. Insect stings inject toxins
into the body that may elicit allergic reactions accompanied by nausea, pain, and swelling.
The bite of the black widow spider is sometimes fatal. Some blood-sucking insects inject
saliva at the point of the bite. Insect saliva may cause swelling and itching, but it may also
carry bacteria or parasites. Bubonic plague, the so-called "black plague" of the middle ages,
which is a bacterial disease, is transmitted by flea bites.

Parasites.

Parasites come into the body through many mechanisms. Inhale the dust of a soiled bed
linen, and you may get pinworms. Take a dip in a lake or river and get schistosomiasis. Get
bitten by a mosquito and get malaria or sleeping sickness. Hug your mother and get follicle
mites. Eat uncooked pork and get trichinosis. Eat food contaminated with fecal matter and
you may get roundworms. Roundworms generally inhabit the intestine, but because of their
complex life cycle, sometimes they end up in other parts of the body, including the
brain. Learn about Hygiene.

Magnetic fields.
We live immersed in the magnetic field of the earth. The human body is generally not
affected and cannot detect this magnetic field. Homing pigeons, however, have been shown
to use the earth's magnetic field as one means for returning home. In principle, however,
the electrical activity of the nervous system could be affected by strong magnetic fields, and
recent experiments suggest that magnetic fields may help to reduce certain kinds of pain.

Gravity.

Our sense of equilibrium in the gravitational field of the earth is provided by the semicircular
canals in the ear. These canals are lined with filaments that are stimulated by calcium
carbonate crystals suspended in a fluid. The rotation of the moon around the earth every 27
1/3 days creates tidal forces that affect many living organisms, but is not known to have a
significant effect on humans. Some fish are known to spawn in the beach at high tide when
the moon is full. The human menstruation cycle of approximately 28 days may be a legacy
of our ancestral origins in the sea. See The Geologic and Biological Timeline of the Earth.

Air pressure.

We can sense changes in air pressure as pain or discomfort in our ears, sinuses or bones.
The nerves surrounding the body cavities that contain enclosed pockets of air detect volume
changes caused by external air pressure.

Endogenic inputs.

Endogenic inputs come from within the body to the brain. When we start exercising, carbon
dioxide builds up in the body. This buildup acts as an endogenic signal for the heart and the
lungs to work harder. When the level of glucose in the blood drops, we get hungry. Hunger,
thirst, pain, fatigue, kinesthesia are all inputs to the brain from the body itself.

Some physiological cycles like menstruation may trigger feelings of fatigue, irritability, and
depression as the hormone levels change. Exercise has been credited with stimulating the
body to generate endorphins that create a feeling of well being. Emotions such as fear
release adrenaline into the bloodstream, which triggers many systemic reactions. Several
studies have found that what you think can affect your health. Constant worry can create
stress that lowers the body's ability to fight diseases, whereas positive thoughts and
laughter can actually improve your health.

Verbal inputs.

Verbal communication might have been included under sounds. However, the effect of
verbal input on the mind is so different from that of the wind blowing or other noises
encountered in nature that it is considered separately. Imagine that your boss calls you to
his office and says something neutral like: "In two weeks we are having a meeting to
discuss the progress of our new project". Your reaction may be one of anticipation or
apathy. Not much is required from you except your participation. However, if the boss says
something negative like: "You made several mistakes in your last report and I am very
dissatisfied with your work". You may become angry or scared, your heart may start racing
and you may want to justify what you did. Words have the power to make you laugh or the
power to make you cry because they are not only sounds. Words have meanings that get to
the root of your emotions.
Since ancient times words have had mystical power because they could represent objects,
feelings, curses, etc. The word "abracadabra" was supposed to have magical powers against
disease or disaster, and sometimes it was carried in an amulet. Prayers were more than just
words; they provided a way of communicating with the deities.

Much can be deduced about the state of mind of a speaker from their speech. The tone of
the voice can convey authority, fear, doubt, and many other different emotions.

Non-verbal sound inputs.

If analyzed carefully, this category could also be grouped under other senses. However,
there are some inputs that connect to the fears or desires deep within our brain and
establish a special kind of non-verbal communication. The snarl of a dog, a cat rubbing
against our legs, a gentle massage, or the wink of an eye are all special kinds of
communication. These are more than just simple sounds or visual or tactile inputs. They are
meaningful messages for the brain.

Blind spot

There are grizzled heroes and sleek assassins in movies who don't have metaphorical
blind spots. They do have literal ones, though. Because of the way the eye is constructed,
every human being does.

For the most part, the human eye gives the brain an accurate picture of what's going on
in the world. There are limitations. Although many birds and insects can see ultraviolet,
and some creatures can see infrared, humans are stuck looking at so-called 'visible' light
only. This cuts down human's view of the world, not letting them see the urine trails left
behind by some mammals, and not letting them fully appreciate the colors of certain
flowers, which have evolved to put on quite a show in ultraviolet while remaining plain
in visible light. The human eye also can't distinguish between polarized and
nonpolarized light, while many cephalopods and some birds can.

Still, the eye sends back signals that let humans navigate through the world pretty
successfully. Many assume that what they see is actually what's out there. That's not
entirely true. Each human eye has a blind spot, and the brain sometimes has to fill in
what is there by looking at the surrounding area.

Light gets into they eye by passing through the pupil. It hits the retina at the back of th
eye. The retina is covered with light-sensing proteins. They relay what they sense to the
optic nerve which carries the information back into the brain. The problem is, the optic
nerve ends in the field of the retina itself. This is a little like having to plug the power
cable for a TV directly into the screen. It creates a dark spot. Most of the time, the other
eye will see what's happening in its partner's blind, but if the blind spots overlap while
looking at a certain object, or if the person is only looking through one eye, the brain just
fills in the spot looking at the surrounding picture.

There's a way to find your blind spot. Cover your left eye and look at the dot on the left in
this image. Be aware of the cross on the right, but don't look at it - just keep your eye on the
dot. Move your face closer to the monitor, and farther away. At some point, you should see
the cross disappear. Stay at that point and close your right eye. Stare at the cross, and you
should see that the dot has disappeared. It doesn't just happen with a white background. Try
the same with colored paper, and your mind will fill in the background color of the paper
when the mark gets in your blind spot. You don't see as much of the world as you think.

Osmosis
and

the Water Potential of Potato Tissue

by

Robert J. Kosinski
Clemson University

Last Update: October 2014

Osmosis has been noticed by biologists since the middle 1700s, and by the 1870s,
careful quantitative observations were being made of it (Baumgarten and Feher,
1998). However, while we can predict it exactly, the cause of osmosis is still in
dispute (Baumgarten and Feher, 1998; Weiss, 1996, p. 216).

Theories on the Cause of Osmosis

The OMP mentioned three theories on the cause of osmosis. A simple and appealing
explanation for osmosis is the concentration of water explanation--water in pure
water is simply more concentrated than water in solutions because the solute has to
take up some room in the solution. According to this idea, water diffuses into a
hyperosmotic solution because it is diffusing down its concentration gradient.
However, a detailed examination reveals problems with this theory. As Weiss (1996)
points out, this predicts water movement in the right direction, but not of the right
magnitude. Water movement in osmosis is faster than diffusion, and seems to be more
like mass water movement caused by a pressure difference (Weiss, 1996, p. 218).
Also, as Salisbury and Ross (1992, p. 39) point out, adding solutes to a solution
decreases the concentration of water in most cases, but in some cases solutions have
a higher concentration of water . The Handbook of Chemistry and Physics has a large
section on solutions of common solutes, and it discloses that a 0.2 M solution of NaCl
has a markedly higher water concentration (995 g/L) than a 0.2 M solution of sucrose
(955 g/L) (Wolf et al., 1982, pp. D261 and D270). Yet our past expereiments have
shown that a potato core loses water to a 0.2 M solution of NaCl but it gains water
from a 0.2 M solution of sucrose. Also, if we compare 0.2 M solutions of sucrose and
glucose, 0.2 M glucose has a higher water concentration than 0.2 M sucrose (976 g/L
vs. 955 g/L) (Wolf et al, 1982, p. D239) . This makes sense because the smaller
glucose molecules take up less room in the solution. However, these two solutions
have exactly the same osmotic potential.

A slightly more complex theory that is often found in general biology books
(including your text, p. 117) is the “bound water” explanation. This says that any
hydrophilic solute (like sucrose or NaCl) will bind up hydrating water and prevent it
from moving freely. Therefore, the side of a semipermeable membrane with pure
water has a higher “free” water concentration than the side with the solute molecules.
According to this explanation, “free” water moves into hypertonic solutions simply
because it is diffusing down its concentration gradient. Although it is popular in
introductory texts, this theory is not even mentioned in several reviews (Baumgarten
and Feher, 1998; Weiss, 1996, pp. 216-222). If the bound water explanation were true,
we would expect that a greater mass of hydrophilic solute would bind more water.
Whether a certain mass of solute is present in a few large molecules or in many small
ones shouldn’t matter. Also, when predicting osmosis, we would have to carefully
consider how hydrophilic the solute is (that is, how many water molecules it binds per
molecule). In fact, the number of molecules present does affect osmosis, and we can
predict osmosis without considering how hydrophilic the solute molecules are.

Another explanation is van't Hoff's Law, or the “number of particles” explanation.

Jacobus H. van’t Hoff (1887) gathered much data on the osmotic potential of many
kinds of solutions. As long as the solution was relatively dilute and the temperature
was constant, he found that the osmotic potential was proportional to
the concentration of solute particles. The size or nature of the solute particles didn’t
matter. So, for example, a small sodium ion would have the same osmotic effect as a
large sucrose molecule, and both of these would be equivalent to a very large starch
molecule (Baumgarten and Feher, 1998). This also means that ionizing substances
like NaCl should have a greater osmotic effect than nonionizing substances like
sucrose because when they ionize, they generate more particles. A measure of
concentration called “osmolality” (the concentration of solute particles in moles/L)
was invented to express this. A 1 molal solution of sucrose would have an osmolality
of 1 osmol/L because each mole of sucrose also produces one mole of solute particles.
However, a completely dissociated 1 molal concentration of NaCl would have an
osmolality of 2 osmol/L because each mole of NaCl ionizes into two moles of
particles. A completely dissociated 1 molal solution of CaCl2 would have an
osmolality of 3 osmol/L because each mole of CaCl2 ionizes into three moles of
particles .

Van't Hoff's Law is an empirical relationship, not a physical description of why

osmosis occurs. Why the number of particles should matter remains unclear. Van't
Hoff himself was only interested in predicting osmosis, and he expressed frustration
with those who wanted him to explain why his law worked (van't Hoff, 1892; quoted
in Weiss, 1996, p. 185). Since even modern experts can't agree why osmosis occurs,
we'll have to follow van't Hoff's example and satisfy ourselves with prediction of
osmosis.

In this exercise, we tested three predictions of Van't Hoff's Law:

The lines describing weight change vs. solution osmolality will the same for the three
solutes;

The no-weight-change osmolality (moles of particles per L at which the line above
crosses the x axis) will be the same for glucose, sucrose, and NaCl;

Van't Hoff's Law will estimate the same water potential for potato cores when using
glucose, sucrose, and NaCl.

The Water Potential of White Potatoes

White potatoes are important food crops in the cooler regions of the world. The tuber,
the edible part of the white potato is a very short and thick, starchy stem, with the
"eyes" being the buds on the stem (Burton, 1989, Chapter 2). White potatoes have
firm tissue and convenient size, so they are favorite subjects for the teaching
laboratory determination of the water potential of plant tissue. 

The "change in weight" method we use in our lab was published in 1935 by Meyer
and Anderson as a modification of a method published by Ursprung (1923). Meyer
and Wallace (1941) employed this method and found that white potato tuber water
potential was between -7.7 and -8.3 bars, with the variance due to the length of time
the sample was tested. Ashby and Wolf (1947) later confirmed these estimates.
Epstein and Grant (1973) used gravimetric methods using sucrose solutions to test two
varieties of white potatoes and found water potentials ranging from -2 to -8 bars.
Ehlenfeldt (1992) used a technique very similar to ours (except that he used sorbitol as
a solute rather than sucrose) and found that the solution with the same water potential
as white potatoes ranged from 0.24 to 0.31 M, corresponding to about -7 bars. Instead
of using a wide range of solutions, as we did, he focused on the point where the line
crosses the x axis by using solutions that ranged from only 0.20 to 0.35 M. Using an
entirely different technique, Gandar and Tanner (1976) used a pressure chamber to
determine water potential in potato tuber cores. Here pressure was applied to the test
specimen until sap just began to wet of the xylem traces at the ends of the cut sample.
They calculated water potential of white potato tubers to be equal to -6 to -7 bars.
Again, extreme variance (up to 5 bars) resulted, due to the moisture of the sample.
Ros Barcelo and Calderon (1994) cited a value of -6.7 bars for white potato tuber
tissue.

Despite the strong agreement in potato water potentials determined by these various
methods, lengthy discussion has centered around the potential sources of error
associated with each method. Meyer and Wallace (1941) suggest that weight change
may not be due to the osmotic movement of water but rather due to the gain or loss of
solutes to or from the soak medium. Bland and Tanner (1985) did a critical
comparison of three methods of determining water potential and found that while two
methods might agree in a certain range of water potentials, in another range they
might diverge widely. Boyer (1969) reviewed numerous methods of water potential
determination.

Growth conditions (especially soil water) cause significant differences in tuber water
potential. As one would expect, drier soils produced drier white potatoes with more
negative water potentials. For example, Burton (1944) found that the water content of
some white potatoes in England increased from 73% to 77% as the rain increased
from 30 mm/month to 70 mm/month. Win et al. (1991) found that white potato tubers
in dry soil in New Jersey increased from a water potential of -4 bars before a
rainstorm to -0.8 bars after the storm. Bland and Tanner (1985) measured the water
potential of white potatoes and found a very wide range, from -0.1 to -0.9 MPa (-1 to -
9 bars), although some stored white potatoes went as low as -15 bars. Bland and
Tanner (1986) traced the drying of stored white potatoes, and found that their water
potentials declined from -3 bars to -5 or -6 bars over the first 5 to 7 weeks of storage.
After 25 weeks of storage, they had gotten down to -7 bars. We have no idea how
long our potatoes were stored. Also, different parts of a tuber might have very
different water potentials. Meissner (1997) found that beet storage organ tissue had a
water potential 5.6 bars lower than the tissue close to the water-conducting vessels
(the xylem). Shibairo et al. (2002) found that weight loss in stored carrots was usually
associated with changes in water potential or osmotic potential under common storage
conditions. 

White potatoes can acclimate to very dry conditions. Leone et al. (1996) found that
the growth of white potato cells in tissue culture would be completely inhibited by
sudden transfer to a solution with an osmotic potential of -23 bars, but the cells could
continue to grow in this solution if they were gradually acclimated to it. Part of the
acclimation was changing the fatty acid composition of the potato cell membranes,
mediated by activation of different genes than in the unstressed cells (Leone et
al., 1996 and 1994). Liu et al. (2006) found that white potato leaf water potentials
declined from -5.3 bars to - 8.5 bars during an experiment testing the effect of water
stress. Vos and Oyarzun (1987) studied the decline in photosynthesis as potato leaves
declined from -6 bars to -11 bars. 

The same general conclusions also apply to sweet potatoes, although all the sweet
potato water potentials tend to be lower. Sung (1985) found that sweet potato leaves
became wilted when the leaf water potential dropped to about -16 bars. Ghuman and
Lal (1983) found that sweet potato leaves in Nigeria had an average water potential of
-9.6 bars. Despite the tough appearance of the tuber, sweet potato is sensitive to water
stress, and the tubers increase their dry matter as water stress gets worse (Ekanayake
and Collins, 2004).

Literature Cited
Ashby, E. and R. Wolf. 1947. A critical examination of the gravimetric method of
determining suction force. Annals of Botany (London) (N.S.) 11(42): 261-268.

Baumgarten, C. M. and J. I. Feher. 1998. Osmosis and the regulation of cell volume.
Pp. 253-292 in N. Sperelakis (Ed.), Cell Physiology Source Book, 2nd Ed. Academic
Press, San Diego, 1095 pp.

Bland, W. L. and C. P. Tanner. 1985. Measurement of the water potential of stored

potato-tubers. Plant Physiology 79: 891-895.

Bland, W. L. and C. B. Tanner. 1986. Potato-tuber water potential components during

storage. American Potato Journal 63(11): 649-653.

Boyer, J. S. 1969. Water status measurements in plants. Annual Reviews of Plant

Physiology 20: 351-364.

Burton, W. G. 1944. The characteristics of certain varieties of potato, with special

reference to their suitability for drying. Annals of Applied Biology 31: 89-96.
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Publishers, Essex, UK, 742 pages.

Ehlenfeldt, M. K. 1992. Evaluation of differential tuber tissue expansion and plant

transpiration as methods for early hollow heart screening. American Potato
Journal 69(9): 537-546.

Ekanayake, I. J. and W. Collins. 2004. Effect of irrigation on sweet potato root

carbohydrates and nitrogenous compounds. Journal of Food, Agriculture and
Environment 2(1): 243-248.

Epstein, E. and W. J. Grant. 1973. Water stress relations of the potato plant under
field conditions. Agronomy Journal 65: 400-404.

Gandar, P. W. and C. B. Tanner. 1976. Potato leaf and tuber water potential
measurements with a pressure chamber. American Potato Journal 53: 1-14.

Ghuman, B. S. and R. Lal. 1983. Mulch and irrigation effects on plant-water relations
and performance of cassava and sweet potato. Field Crop Research7(1): 13-29.

Leone, A., A. Costa, S. Grillo, M. Tucci, I. Horvath and L. Vigh. 1996. Acclimation
to low water potential determines changes in membrane fatty acid composition and
fluidity in potato cells. Plant, Cell & Environment 19(9): 1103-1109.

Leone A., A. Costa, M. Tucci and S. Grillo. 1994. Comparative analysis of short- and
long-term changes in gene expression caused by low water potential in potato
(Solanum tuberosum) cell-suspension cultures. Plant Physiology 106(2): 703-712.

Liu, F., A. Shahnazari, M. N. Andersen, S. E. Jacobsen and C. R. Jensen. 2006.

Effects of deficit irrigation (DI) and partial root drying (PRD) on gas exchange,
biomass partitioning, and water use efficiency in potato. Scientia
Horticulturae 109(2): 113-117.

Meissner, S. T. 1997. Water potential measurements of the storage parenchyma tissue

and xylem solution of red beets (Beta vulgaris) suggests apoplastic discontinuity
between the conducting xylem and sink areas. The American Journal of Botany 84(6):
S150.

Meyer, B. S. and D. B. Anderson. 1935. Laboratory Plant Physiology. 107 pp.

Edwards Bros., Ann Arbor.
Meyer, B. S. and A. M. Wallace. 1941. A comparison of two methods of determining
the diffusion pressure deficit of potato tuber tissues. American Journal of Botany 28:
838-843.

Ros Barcelo, A. and A. A. Calderon. 1994. Measuring water conductivity coefficients

in plant tissues. Journal of Biological Education 28(2): 83-85.

Salisbury, F. B. and C. W. Ross. 1992. Plant Physiology, 4th Ed. Wadsworth

Publishing Co., Belmont, CA. 682 pp.

Shibairo, S.I., M.K. Upadhyaya, and P.M.A. Toivonen. 2002. Changes in water
potential, osmotic potential, and tissue electrolyte leakage during mass loss in carrots
stored under different conditions. Scientia Horticulturae 95(1): 13-22.

Sung, J.-M. 1985. Studies on physiological response to water stress in sweetpotato. II.
Osmotic adjustment in sweetpotato leaves. Journal of the Agricultural Association of
China 129: 50-55.

Ursprung, A. 1923. Zur Kenntnis der Saugkraft. VII. Eine neue vereinfachte Methode
zur Messung der Saugkraft. Ber. Deutsch Bot. Ges. 41: 338-343.

van't Hoff, J. H. 1887. Die Rolle des osmotischen Drukes in der Analogie zwischen
Losungen und Gasen. Z. Physik. Chemie 1: 481-493.

van't Hoff, J. H. 1892. Zur Theorie der Losungen. Z. Physik. Chemie 9: 477.

Vos, J. and P. J. Oyarzun. 1987. Photosynthesis and stomatal conductance of potato

leaves--effects of leaf age, irradiance, and leaf water potential.Photosynthesis
Research 11: 253-264.

Weiss, T. F. 1996. Cellular Biophysics, Vol. 1: Transport. MIT Press, Cambridge,

MA. 693 pp.

Win, K., G. A. Berkowitz and M. Henninger. 1991. Antitranspirant-induced increases

in leaf water potential increase tuber calcium and decrease tuber necrosis in water-
stressed potato plants. Plant Physiology 96: 116-120.

Wolf, A. V., M. G. Brown and P. G. Prentiss. 1982. Concentrative properties of

aqueous solutions: Conversion tables. Pp. D227-D276 in R. C. Weast and M. J.
Astle, CRC Handbook of Chemistry and Physics, CRC Press, Boca Raton, Florida.
 Coursewide Data

These data were based on 33 water potential values estimated with NaCl, 32 with sucrose, and 34
with glucose. The experiment was done during the week of September 29, 2014. 
 

Table 1. Average percent change in mass of white potato cores during a 1.5-hour
incubation in several NaCl, sucrose, and glucose solutions. 
Molality NaCl Sucrose Glucose

0.0 +9.30% +9.08% +10.73%

0.1 +2.87% +5.16% +5.69%

0.2 -7.88% +2.48% +3.33%

0.3 -17.40% -3.02% -1.44%

0.4 -23.35% -10.05% -6.46%

0.5 -26.96% -14.93% -12.82%

You should use these data in order to make an Excel graph (Figure 1) in your report.
This graph should plot the percent weight change (y axis) vs. osmolality of the soak
solution (x axis). Each graph should have a line for the 2014 coursewide NaCl,
glucose, and sucrose averages above plus a fourth line for your lab group's data (either
NaCl, glucose, or sucrose), as was illustrated in the OMP. These three lines should be
distinctive (e.g., points with different shapes and dotted vs. solid lines). Use the
techniques you practiced in the data presentation exercise the second week of lab.
This implies that graphs will not have color, will use only circles, squares and
triangles (black or white) for the points, and that the lines will be solid black or
dashed in various ways.

All of these x axis variables can be derived from the molality data above. Remember,
osmolality = molality for sucrose and glucose (because sugars do not ionize), and
osmolality = 1.8x molality for NaCl (because NaCl ionizes into two particles). The
molecular mass of NaCl is 58.4; the molecular mass of sucrose is 342.3; the molecular
mass of glucose is 180.2.
Then you will need a table. Remember, the different solutes are the treatments in this
case, and we are testing to see if the treatments are the same. This single table should
summarize the chi-square results below for the following questions:

Are the potato water potentials estimated from the NaCl, glucose, and sucrose data the
same?

Do potato cores have no weight change in the same osmolalities of NaCl, glucose, and
sucrose?

You will need to summarize these data in a table that shows the means of each no-
weight-change variable for NaCl, glucose, and sucrose, the chi-square for the test of
equality of NaCl, glucose, and sucrose for each variable, and the P-value for each
variable's test. Note that you do not have to include the above- and below-median data
in the tables below. It might be a good idea to devote one line of your table to each
variable.

Table 2. Potato water potentials (bars) estimated by van't Hoff's Law for NaCl, sucrose and
glucose, and the results of a chi-square median test comparison of the water potential estimates
by the three solutes.

Means Below Above

Median Median
NaCl -5.67 12  21

Sucrose -5.66 13.5 18.5

Glucose -5.92 24 10

Chi-square = 9.00; P value = 0.011

In interpreting the table above, remember that -5.92 bars is below -5.66 bars.

Table 3. Osmolalities (moles of solute particles/kg of water) of NaCl, sucrose, and glucose that
produced zero weight change in potato cores, and the results of a chi-square median comparison
of the no-weight-change osmolality for the three solutes.

Means Below Above

Median Median
NaCl 0.191 25.0  8.0

Sucrose 0.226 13.5 18.5

Glucose 0.167 11.0 23.0 

Chi-square = 13.774; P value = 0.001