User:Pflanzenliebhaber749/sandbox

General Article Review

Overall article could be re-written for better grammar, and the table of contents could be expanded in terms of how certain factors tie into turgor pressure within plants, as measurements of turgor pressure in general (including units), other applications of turgor pressure outside of plant physiology, how turgor pressure is measured within plants. Mostly found some basic biology sites which I used to help understand the basic mechanisms within this field, but things such as journals and research papers will be used to explain the more complex areas/mechanisms of turgor pressure within plants. I don't want this article to rely too heavily on turgor pressure within plants themselves, so I will have to find more sources about turgor pressure in outside applications, protists, the reasons why it isn't observed in animal cells, and observations of it in bacterial and fungal cells. Hydrostatic pressure can also be explained somewhat, but there is already a Wikipedia page on it. I can link this in with a section where I can briefly explain the discrepancies between the two though

Turgor Pressure Article Draft (Lead)

Turgor pressure is the force within a cell that pushes the plasma membrane against the cell wall. It is also called Hydrostatic pressure, and more intricately defined as the pressure exerted by a fluid, measured at a certain point within itself when at equilibrium. Generally, turgor is caused by the osmotic flow of water, and occurs in plants, fungi and bacteria. The phenomenon is also observed in protists that have cell walls. This system is not seen in animal cells, seeing how the absence of a cell wall would cause the cell to lyse when under too much pressure. The pressure exerted by the osmotic flow of water is also known as turgidity. It is caused by the osmotic flow of water through a semipermeable membrane. Osmotic flow of water through a semipermeable membrane is when the water travels from an area with a high-solute concentration, to one with the lower-solute concentration. In plants, this entails the water moving from the low concentration solute outside the cell, into the cell’s vacuole.

Mechanisms

Osmosis is the process in which water flows from an area with a high solute concentration, to an adjacent area with a lower solute concentration until equilibrium between the two areas is reached. All cells are surrounded by a lipid bilayer cell membrane which permits the flow of water in and out of the cell and limits the flow of solutes. When in a hypotonic solution, water flows into the membrane and increases the cell’s volume, and when in an isotonic solution, water flows in and out of the cell at an equal rate.

Turgidity is the point at which the cell’s membrane pushes against the cell wall, which is when turgor pressure is high. When the cell membrane has low turgor pressure it is then flaccid. In plants we see this as wilted anatomical structures. This is more specifically known as plasmolysis.

The volume and geometry of the cell affects turgor pressure values and how turgor pressure’s effect on cell wall plasticity. Studies have shown how smaller cells experiences a stronger elastic change when compared to larger cells.

Turgor Pressure in Plants

Turgor pressure within cells is regulated by osmosis and also causes the cell wall to expand during growth. Along with size, rigidity of the cell is also caused by turgor pressure; and a lower pressure results in a wilted cell or plant organism (ie. leaf, stalk). One mechanism in plants that regulate turgor pressure include semi-permeable membranes, which only allow some solutes to travel in or out of the cell, which can also and somewhat maintain a certain minimum of turgor pressure. Other mechanisms include transpiration, which results in water loss and a decrease in turgidity in cells. Turgor pressure is also a large factor for nutrient transport throughout the plant. Cells of the same organism can have differing turgor pressure throughout its structure, which is responsible for certain functions of structures. In higher plants, turgor pressure is responsible for apical growth of things such as root tips and pollen tubes.

Dispersal

Transport proteins that pump solutes into the cell can be regulated by cell turgor pressure. Lower values allow for an increase in pumping of solutes which in turn increases osmotic pressure. This function is important as a plant response under drought conditions (seeing as turgor pressure is maintained), and for cells which much accumulate solutes (i.e developing fruits).

Flowering and Reproductive Organs

It has been recorded that petal of Gentiana kochiana and Kalanchoe blossfeldiana bloom via volatile turgor pressure of cells on the plant's adaxial surface. During processes like anther dehiscence, it has been observed that drying endothecium cells causes an outward bending force leading to the release of pollen. This means that lower turgor pressures are observed in these structures, seeing as they're dehydrated. Pollen tubes are cells which elongate when pollen lands on the stigma, at the carpal tip. These cells grow rather quickly due to increasing turgor pressure. These cells undergo tip growth, and Lilies can have a turgor pressure of 0-21 MPa when growing.

Mature squirting cucumber

Seed dispersal

In fruits such as Impatiens parviflora, Oxalis acetosella and Ecballium elaterium, turgor pressure is the method by which seeds are dispersed. In Ecballium elaterium, or squirting cucumber, turgor pressure builds up in the fruit to the point that it aggressively detaches from the stalk, and seeds and water are squirted everywhere as the fruit falls to the ground. Turgor pressure within the fruit ranges from .003-1.0 MPa.

Tree roots begin to split a boulder.

Growth

Turgor pressure actions on extensible cell walls is usually said to be the driving force of growth within the cell. An increase of turgor pressure causes expansion of cells and extension of apical cells, pollen tubes and in structures such as root tips. Cell expansion and an increase in turgor pressure are due to inward diffusion of water into the cell, and turgor pressure increases due to the increasing volume of vacuolar sap. A growing root cell's turgor pressure can bee up to 6 bars, over three times that of a car tire! Epidermal cells in a leaf can have pressures ranging from 15-20 bars. Seeing that plants operate at such high pressures, it can explain why trees can grow through asphalt and other hard substances.

Turgidity

This is observed in a cell when the cell membrane is pushed against the cell wall. In some plants, their cell walls loosen at a quicker than water can cross the membrane, which results in a cell with lower turgor pressure.

Stomata

Open stomata on the left and closed stomata on the right

Turgor pressure within the stomata regulates when the stomata can open and close, which has a play on transpiration rates of the plant. This is also important because this function regulates water loss within the plant. When turgor pressure is low, the stomata closes. Lower turgor pressure can mean that the cell has a low water concentration and closing the stomata would help conserve water. High turgor pressure keeps the stomata open for gas exchanges necessary for photosynthesis.

Mimosa pudica

Mimosa pudica

It has been concluded that loss of turgor pressure within the leaves of mimosa pudica is responsible for the reaction the plant has when touched. Other factors such as change in osmotic pressure, protoplasmic contraction and increase in cellular permeability have been observed as well.  It has been recorded that turgor pressure is to be different between upper and lower pulvinar cells of the plant, and the movement of potassium and calcium ions throughout the cells which causes an increase in turgor pressure. When touched, the pulvinus is activated and exudes contractile proteins, which in turn increases turgor pressure and closes the leaves of the plant.

Function in Other Taxa

As earlier stated, turgor pressure can be found in other organisms besides plants, and can play a large role in the development, movement and nature of said organisms.

Fungi

Shaggy Ink Caps busting through asphalt due to high turgor pressures.

In fungi, turgor pressure has be observed as a large factor in substrate penetration. In species such as Saprolegnia ferax, Magnaporthe grisea, and Aspergillus oryzae, immense turgor pressures have been observed in hyphae which can penetrate substances like plant cells, synthetic materials such as polyvinyl chloride, asphalt and plastics. In observations of this phenomenon, it is noted that invasive hyphal growth is due to turgor pressure, along with the coenzymes secreted by the fungi to invade said substrates. Hyphal growth is directly related to turgor pressure, and growth slows as turgor pressure decreases. In Magnaporthe grisea, pressures of up to 80 bars have been observed.

Protists

Some protists do not have cell walls and cannot experience turgor pressure. These few protists are ones which use their contractile vacuole which regulates the quantity of water within the cell. Protist cells avoid lysing in solutions by utilizing a vacuole which pumps water out of the cell to maintain osmotic equilibrium.

Animals

Turgor pressure is not observed in animal cells because they lack a cell wall. In organisms with cell walls, the cell wall prevents the cell from lysing from high pressure values.

Diatoms

In Diatoms, the Heterokontophyta have polyphyletic turgor-resistant cell walls. Throughout these organisms’ life-cycle, carefully controlled turgor pressure is responsible for cell expansion and for the release of sperm, but not for seta growth.

Cyanobacteria

Gas-vacuolate cyanobacterium are the ones generally responsible for water-blooms. They have the ability to float due to the accumulation of gases within their vacuole, and the role of turgor pressure and its effect on the capacity of these vacuoles has been observed in varying scientific papers. It is noted that the higher the turgor pressure, the lower the capacity of the gas-vacuoles in different cyanobacterium. Experiments used to correlate osmosis and turgor pressure in prokaryotes have been used to show how diffusion of solutes into the cell have a play on turgor pressure within the cell.

Measurements

When measuring turgor pressure in plants, many things have to be taken into account. It is generally stated that fully turgid cells’ turgor pressure value is equal to the osmotic potential of the cell, and that flaccid cells’ values are at or near zero. When measuring turgor pressure factors such as osmotic pressure, and  total water potential. Other cellular mechanisms taken into consideration include the protoplast, solutes within the protoplast (solute potential), transpiration rates of the plant, and tension of the cell’s wall, just to name a few. Measuring turgor pressures within plants can have their own limitations depending on the methods used .Of course, there are many methods of measuring turgor pressure in plants and other organisms, of which will be explored and explained below. Not all methods can be used for ‘all’ organisms due to their sixes, or something as simple as the nature of the organism (i.e a diatom won’t have the same properties of a plant, which would place constrictions on what could be used to infer turgor pressure).

Units

Units used to measure turgor pressure are independent from the measures used to infer its values. Common units used are bars (15 lbs per square inch) MPa or newtons/millimeter squared. One bar is equal to 10 MPa.

Methods

Water Potential Equation

Turgor pressure can be deduced when Ψw (total water potential) and Ψs  (osmotic potential) are known in a water potential equation. Water potential equations are used to measure total water potential of a plant by using variables such as matric potential, osmotic potential, pressure potential, gravitational effect and of course, turgor pressure. When the difference is taken between Ψw and Ψs, we are given the value for turgor pressure. When using this method, factors like Ψg are considered negligible since is doesn’t  have that great of an effect on measurements in short plants. Matric potential,Ψm, is also ignored because it’s values are always negative or very close to zero.

Pressure-Bomb Technique

Pressure bomb diagram

The pressure bomb was first developed by P. F. Scholander and his colleagues in 1965, who used the first pressure bomb to speculate water movement through plants. The first commercially available chamber was made 2 years later. In the 1970’s, this instrument was widely used in laboratories which studied plant physiology. The instrument is used to measure turgor pressure by placing a leaf (with its stem attached) into a closed chamber, and pressurized gas is added in increments. Measurements are taken when xylem solution  appears out of the cut surface, and is at the point where the xylem doesn’t accumulate or retreat back into the cut surface.

Atomic-Force Microscope

Atomic force microscopes are a type of scanning probe microscope (SPM), which are used to measure properties within an area via scanning. Small probes are introduced to an area of interest and can find measurements for things such as height, width and friction. The Atomic force microscope is one that utilizes a spring placed within the probe, which measures values via displacement of the probe. This is used when measuring turgor pressure of organisms. When using this method, supplemental information such as single force depth curves, continuum mechanic equations and cell geometries can be used to quantify turgor pressures within a given area (usually a cell).

Pressure Probe

This was originally used to measure individual algal cells, and is used on specimen with larger cells. These are usually used on higher plant tissues, and aren’t generally used when measuring turgor pressure. Hϋsken and Zimmerman were scientists that helped to improve this method of measurement, which allowed for a larger amount of researchers to use this method to measure things such as turgor pressure. When finding turgor pressure pressure probes measure by displacement, like the atomic force microscopes do. A glass microcapillary tube is inserted into the cell and whatever the cell exudes into the tube is observed by a microscope, and an attached device measures how much pressure is required to push that excretion back into the cell.

For smaller cells, things such as micromanipulation probes could be used to more accurately quantify measurements. In an 2006 experiment by Wang, Thomas, Pritchard and Hukin, single tomato cells were compressed between a micromanipulation probe and glass, then a pressure probe’s microcapillary was introduced to the cell and used to find turgor pressure(insert citation).

Theoretical Speculations

'Negative' Turgor Pressure

It has been observed that the value of  Ψw  decreases was a cell becomes more dehydrated, but scientists have speculated whether this value will just continue to decrease and never fall below zero, or if the value can be less than zero ( negative). There have been studies which show that negative cell pressures can exist in xerophytic plants, but a paper by M. T. Tyree, explores the notion of  if this is actually possible, or just a conclusion based on misinterpreted data. In this, he concludes that by mis-categorizing “bound” and “free” water in a cell, researchers that claimed to have found negative turgor pressure values are incorrect. By analyzing isotherms of apoplastic and symplastic water, he shows that negative turgor pressures cannot be present within arid plants due to the net water loss of the specimen during droughts. Of course, this is just his analyzation and interpretation of data, and negative values for  Ψp are still used within the scientific community.

Tip Growth in Higher Plants

An hypothesis formed by scientist M. Harold and his colleagues suggests that tip growth in higher plants is amoeboid in nature, not caused by turgor pressure as is widely believed. The hypothesis proposes that growth in higher plants are amoeboid, meaning that extension is caused by the actin cytoskeleton in these plant cells. Regulation of cells growth is implied to be caused by cytoplasmic microtubules which control the orientation of cellulose fibrils which are deposited into the adjacent cell wall, resulting in growth. In plants, the cells are surrounded by cell walls and filamentous proteins which retain and adjust plant cell’s growth and shape. As explained in the paper, lower plants grow through apical growth, which differs since the cell wall only expands only on one end of the cell.

History of the interpretation of Turgor Pressure and disambiguities

It is hard to exactly pinpoint when speculations of turgor pressure began.

It may also be easy for one to confuse turgor pressure with other processes such as osmosis, hydrostatic pressures and other phenomena within fluid mechanics or physiological sciences.