What is Turgor Pressure?
- Turgor pressure, scientifically defined, is the internal force exerted by water against the plasma membrane and the subsequent counterforce of the cell wall. Originating from the Latin term “turgidus,” which means swollen or distended, this pressure is pivotal in maintaining the structural integrity of certain cells, especially in plants, fungi, and bacteria.
- This pressure is a direct consequence of the osmotic flow of water. Osmosis, a fundamental biological process, involves the movement of water molecules through a selectively permeable membrane from a region of low solute concentration to one of higher concentration. In the context of plant cells, water tends to move from the external environment with a lower solute concentration into the cell’s vacuole, which typically has a higher solute concentration.
- The term “turgidity” is used to describe the state of a cell when it is swollen due to the osmotic flow of water. It’s worth noting that while turgor pressure is a common phenomenon in organisms with cell walls, such as plants, fungi, and certain protists, it is absent in animal cells. The lack of a protective cell wall in animal cells means that excessive internal pressure would lead to cell lysis, or rupture.
- The structural rigidity in plants, fungi, and certain bacteria is attributed to turgor pressure. These organisms secrete extracellular molecules that coalesce to form a robust cell wall. As water fills these cells, it exerts pressure against both the plasma membrane and the cell wall, leading to turgor pressure. This pressure not only prevents the cell from bursting in hypotonic environments but also plays a crucial role in the morphology and movement of multicellular organisms.
- For instance, plants have specialized vacuoles that regulate turgor pressure. These vacuoles, being hypertonic relative to the cytoplasm, draw water out of it, ensuring a consistent cytoplasmic concentration. As the vacuole accumulates water, the resulting turgor pressure pushes against the cell boundaries. This mechanism allows plants to maintain their shape, and by modulating turgor pressure in specific cells, plants can even exhibit movements, such as turning their leaves towards sunlight.
- To sustain optimal turgor pressure, plants and fungi ensure that their internal cellular environment remains hypotonic. This concentration gradient facilitates the upward movement of water, nourishing all cells. Once a balance is achieved, excess water is typically evaporated from the organism’s surface or leaves, ensuring a continuous nutrient flow from the roots. In contrast, animal cells, lacking a cell wall, employ mechanisms to maintain isotonic conditions, preventing the onset of turgor pressure and potential cell rupture.
- In conclusion, turgor pressure is a fundamental biological concept, playing a pivotal role in the structural and functional aspects of various organisms. Its understanding is crucial in the realms of cellular biology and plant physiology.
Definition of Turgor Pressure
Turgor pressure is the internal force exerted by water against a cell’s plasma membrane and its supporting cell wall, primarily observed in plants, fungi, and bacteria, resulting from the osmotic flow of water into the cell.
Mechanism of Turgor Pressure
Turgor pressure, a fundamental concept in cellular biology, is primarily observed in cells with a cell wall, such as those in plants, fungi, and certain bacteria. The mechanism of turgor pressure can be understood by examining its behavior in two distinct cellular states: the flaccid cell and the turgid cell.
A. Flaccid Cell: Often referred to as a shrink cell, the flaccid cell exhibits a turgor pressure of zero. The relationship between the diffusion pressure deficit (DPD) and turgor pressure (TP) in this state is defined by the equation:
DPD (diffusion pressure deficit)=OP (osmotic pressure)−TP (turgor pressure)DPD (diffusion pressure deficit)=OP (osmotic pressure)−TP (turgor pressure)
Given that the turgor pressure in a flaccid cell is zero, the DPD equates to the osmotic pressure. This implies that the osmotic potential of the flaccid cell aligns with that of the surrounding environment.
B. Turgid Cell: In the turgid state, the cell’s turgor pressure matches the osmotic pressure, rendering the DPD value to zero. This condition signifies that the cell is in an equilibrium state with its external environment, with no net movement of water in or out of the cell.
It’s crucial to note that the mechanism of turgor pressure is absent in animal cells. Unlike plant cells, animal cells are devoid of a cell wall and possess only a plasma membrane. In the absence of a protective cell wall, excessive internal pressure could lead to cell rupture or lysis. Consequently, turgor pressure is not a feature of animal cells.
In summary, turgor pressure is a result of the osmotic balance between the cell and its external environment, and its mechanism is intricately linked to the presence of a cell wall and the osmotic potential of the cell.
Turgor Pressure and Osmosis
In the realm of cellular biology, the movement of water molecules across biological membranes plays a pivotal role in maintaining cellular homeostasis. This passive transport process, termed osmosis, involves the diffusion of water molecules from a region of low solute concentration to one of higher solute concentration. Alternatively, osmosis can be defined as the net movement of water molecules across a membrane from an area with a higher water potential to one of lower water potential.
Turgor pressure, a direct consequence of osmotic water movement, is particularly significant in plant cells. When there is a net positive influx of water into the cell, it results in turgidity. Unlike animal cells, where excessive osmotic influx can lead to cell lysis or bursting, plant cells are safeguarded by their rigid cell walls. These walls prevent the cells from bursting and, in fact, rely on turgor pressure to maintain their structural integrity and rigidity.
Mechanism of Turgor Pressure Maintenance in Plants: Central to the maintenance of turgor pressure in plant cells is the vacuole, an organelle predominantly filled with water. Positioned between the cell membranes, the vacuole stores water, exerting pressure against the cell wall. This pressure, known as turgor pressure, is essential for the plant’s structural support. In the absence of adequate turgor pressure, plants exhibit a wilted appearance.
Often referred to as hydrostatic pressure, turgor pressure is vital for plants to maintain their structural rigidity. This pressure is a manifestation of the fluid’s force pushing against the cell wall.
Significance of Vacuoles in Plant Cells: Vacuoles, prominent organelles in plant cells, are instrumental in upholding the majority of the cell’s turgor pressure. Apart from their role in turgor maintenance, vacuoles assist in detoxifying the cell by sequestering harmful waste products. These organelles, encapsulated by a single membrane, also play a role in regulating the intracellular pH by housing small compounds that modulate the acidic environment. Furthermore, the vacuolar membrane serves multifaceted functions, including providing shape to the cell.
In conclusion, the interplay between osmosis and turgor pressure is fundamental to the survival and functionality of plant cells. The vacuole, as a central organelle, plays a crucial role in this dynamic equilibrium, ensuring the plant’s structural and physiological well-being.
Turgor Pressure in Plants
Turgor pressure is a fundamental physiological phenomenon observed in plant cells, playing a pivotal role in maintaining the structural integrity and optimal functionality of plants. This pressure arises due to the osmotic flow of water into the cell, exerting force against the cell wall.
- Role in Plant Rigidity: When observing two contrasting plant systems, system A and system B, system A represents cells that are densely packed, akin to a brick wall, while system B signifies cells that are loosely arranged. In system A, the cells exhibit a higher water concentration, leading to elevated turgor pressure compared to the flaccid cells of system B with reduced turgor pressure. This turgor pressure is instrumental in ensuring that plants remain upright and rigid. A loss of this pressure results in the cell becoming flaccid, leading to plant wilting.
- Cellular Mechanisms for Turgor Regulation: Plant cells are equipped with specialized structures that facilitate the regulation of internal turgor pressure. The presence of a cell wall, primarily composed of cellulose, provides the necessary rigidity to the plant cells. This wall prevents the cell from bursting even when there’s a substantial influx of water. Some plant cells possess a dual-layered cell wall: a primary layer and a secondary layer enriched with lignin, which imparts waterproofing properties.
- Vacuoles and Osmoregulation: Central to the cell’s osmoregulatory processes is the vacuole, a large vesicle that dominates the cell’s interior. The vacuole plays a pivotal role in maintaining turgor pressure by regulating the osmotic flow of water. By adjusting the concentration of solutes within the cell, the vacuole ensures a hypertonic intracellular environment, promoting water influx and resulting in increased turgor pressure.
- Turgor Pressure and Stomatal Function: Stomata, tiny pores present on plant surfaces, primarily leaves, facilitate gas exchange. The opening and closing of these stomata are regulated by guard cells, which utilize turgor pressure. When guard cells are turgid due to increased internal water content, stomata open, allowing for the exchange of gases like carbon dioxide and oxygen, essential for photosynthesis.
- Implications of Negative Turgor Pressure: While most plant cells exhibit positive turgor pressure, the xylem tissue, responsible for transporting water and nutrients, experiences negative turgor pressure during transpiration. As plants lose water through evaporation, the resultant high surface tension in the xylem facilitates water transport from the roots to the plant’s upper regions.
In conclusion, turgor pressure is indispensable for plants, influencing their structure, growth, and various physiological processes. Whether it’s providing rigidity, facilitating gas exchange, or ensuring nutrient transport, turgor pressure remains central to plant health and vitality.
Function of Turgor Pressure in Plants
- Maintenance of Structural Rigidity: Turgor pressure plays a pivotal role in ensuring the structural integrity of terrestrial plants. By providing the necessary turgidity, it enables plants to maintain an upright posture, counteracting gravitational forces. This rigidity is essential for plants to position themselves optimally towards light sources, facilitating efficient photosynthesis.
- Regulation of Stomatal Activities: Guard cells utilize turgor pressure to orchestrate the opening and closing of stomata, microscopic pores on plant surfaces. These stomata are integral to various physiological processes, including transpiration, water movement, and photosynthesis. The regulation of stomatal activities is directly influenced by the turgor pressure within the guard cells.
- Induction of Nastic Movements: Certain plants exhibit nyctinastic movements, where they adopt specific postures during the day and night. By day, these plants remain erect, maximizing light absorption for photosynthesis. As night approaches, they exhibit a drooping posture, often closing their leaves and flowers. This movement is attributed to the turgor pressure changes within the pulvinar cells located at the leaf base or petiole apex. A classic example of this phenomenon is observed in Mimosa pudica, where leaves respond to touch by altering turgor pressure within pulvinar cells.
- Facilitation of Apical Growth: Turgor pressure is instrumental in plant growth, particularly at the apical regions. As the cell wall expands under the influence of turgor pressure, it promotes the elongation of root tips, contributing to the overall growth of the plant.
- Mechanism for Seed Dispersal: Certain plants, such as Ecballium elaterium (squirting cucumber), harness turgor pressure for seed dispersal. The accumulated pressure within the fruit leads to its detachment from the stalk. Upon detachment, the internal pressure propels the seeds and internal water out, aiding in seed dispersal. The internal pressure within these fruits can range from 0.003 to 1.0 MPs, highlighting the significant force exerted by turgor pressure.
In summary, turgor pressure is not merely a passive physiological phenomenon in plants. It actively participates in a myriad of essential functions, from maintaining structural rigidity to facilitating growth and reproduction. Its multifaceted roles underscore its significance in plant biology and ecology.
Turgor Pressure in Non-Plant Organisms
1. Bacteria and Turgor Regulation: Bacteria, particularly those possessing cell walls, exhibit remarkable resilience against cell lysis due to their robust cell walls primarily composed of peptidoglycan. While distinct from plant cell walls, bacterial walls serve a similar protective function, resisting potential damage during excessive water influx. Bacteria employ various strategies to modulate turgor pressure:
- Cell Wall Modification: In hypertonic environments, bacteria enhance their resilience by adding more peptidoglycan layers, preventing excessive water loss. Conversely, in hypotonic conditions, they degrade parts of their cell wall to allow easier water entry.
- Osmoprotectant Production: Bacteria synthesize osmoprotectants or osmolytes, organic compounds that elevate the solute concentration in their cytoplasm, counteracting external osmotic pressures.
- Polysaccharide Secretion: Certain bacteria secrete polysaccharides that form a protective coat around the cell, preventing water loss and aiding in turgor pressure maintenance under adverse conditions.
2. Protists and Osmoregulation: Plant-like protists, such as algae, possess cell walls with cellulose, akin to plants. However, variations exist; for instance, diatoms have silica-enriched cell walls, while calcareous algae contain calcium carbonate. Algae regulate turgor pressure through:
- Ion Transport Mechanisms: Algae utilize active transport systems and ion channels to manage ion concentrations, influencing osmotic balance.
- Osmolyte Production: Similar to bacteria, algae produce osmolytes to adjust their internal solute concentrations.
- Vacuolar Systems: Algal cells contain vacuoles that play a role in osmoregulation, ensuring optimal turgor pressure.
3. Fungi and Turgor Maintenance: Fungi, with cell walls predominantly composed of chitin, a complex sugar, exhibit a robust barrier against water loss, especially in hypertonic environments. Turgor pressure in fungi is crucial for various physiological processes:
- Cell Shape Maintenance: Turgor pressure ensures that fungal cells retain their characteristic shape.
- Nutrient Uptake: Proper turgor pressure facilitates efficient nutrient absorption.
- Reproduction: Turgor pressure aids in the effective discharge of spores during fungal reproduction.
- Host Penetration: Fungi utilize turgor pressure to penetrate host cells or substrates, a critical aspect of their parasitic or symbiotic relationships.
In conclusion, while turgor pressure is often associated with plants, its significance extends to various non-plant organisms, including bacteria, protists, and fungi. These organisms have evolved unique mechanisms to regulate and utilize turgor pressure, underscoring its universal importance in cellular biology.
Measurement Methods and Unit of Turgor Pressure
Turgor pressure, a fundamental aspect of plant physiology, is the force exerted by water against the cell wall, providing structural support to the plant. Accurately measuring this pressure is crucial for understanding various cellular processes and plant responses to environmental conditions. However, the measurement of turgor pressure is intricate, necessitating consideration of various cellular components and mechanisms.
Units of Measurement: Turgor pressure is quantified using specific units, with the most common being bars, MPa (megapascals), and newtons per square meter. Notably, 1 bar is equivalent to 0.1 MPa.
Methods to Measure Turgor Pressure:
- Water Potential Equation: By understanding the total water potential (Ψw) and the osmotic potential (Ψs), one can deduce turgor pressure. The equation considers various factors, including matric potential, osmotic potential, pressure potential, and gravitational effects. Typically, the effects of gravity and matric potential are negligible, allowing for a more straightforward calculation.
- Pressure-bomb Technique: Introduced by Scholander and later reviewed by Tyree and Hammel, this technique involves placing a leaf (with an attached stem) inside a chamber. The chamber is then pressurized incrementally until xylem sap emerges from the cut surface. The point at which the sap neither accumulates nor retreats determines the turgor pressure.
- Atomic Force Microscope (AFM): Operating under the principles of scanning probe microscopy, AFM uses minute probes to measure displacement, thereby determining turgor pressure. This method requires additional data, such as continuum mechanic equations and cell geometries, to provide accurate measurements.
- Pressure Probe: Initially designed for algal cells, this device has since been adapted for larger specimens, predominantly higher plant tissues. The method involves inserting a glass micro-capillary tube into the cell and observing the cellular exudate. An attached instrument then calculates the pressure needed to reintroduce the exudate into the cell.
- Micro-manipulation Probe: Ideal for smaller cells, this probe provides precise measurements. In a study conducted by Weber and Smith, individual tomato cells were compressed using a micro-manipulation probe, enabling the pressure probe to determine the cell’s turgor pressure.
In conclusion, while the measurement of turgor pressure is complex, various innovative techniques have been developed to ensure accuracy. Each method, with its unique approach, contributes to a comprehensive understanding of this vital cellular phenomenon.
Theoretical speculations of Turgor Pressure
Turgor pressure, a pivotal concept in plant physiology, has been the subject of various theoretical speculations. These speculations delve into the nuances of turgor pressure and its implications in plant cellular mechanisms.
- Negative Turgor Pressure: A perplexing observation in plant physiology is the decreasing value of water potential (Ψw) as cells undergo dehydration. This has led to debates regarding the possibility of Ψw values becoming negative. While certain studies have indicated the existence of negative cell pressures in xerophytic plants, M. T. Tyree critically examined this claim. Tyree posited that the reported negative turgor pressures were erroneous, stemming from a misinterpretation of the distinction between “bound” and “free” water within a cell. Through a meticulous analysis of apoplastic and symplastic water isotherms, Tyree demonstrated that negative turgor pressures are implausible in arid plants, primarily due to the net water loss these plants experience during drought conditions. Notwithstanding Tyree’s analysis, the notion of negative turgor pressures persists in some scientific circles.
- Tip Growth in Higher Plants: The conventional understanding attributes tip growth in higher plants to turgor pressure. However, a hypothesis presented by M. Harold and his team challenges this belief. They propose that tip growth in higher plants exhibits amoebic characteristics and is not driven by turgor pressure. Instead, they suggest that the actin cytoskeleton within plant cells is responsible for extension. This theory emphasizes the role of cytoplasmic micro-tubules in regulating cell growth. These micro-tubules determine the orientation of cellulose fibrils, which are subsequently incorporated into the neighboring cell wall, culminating in growth. Furthermore, the unique structure of plant cells, encompassed by cell walls and filamentous proteins, plays a crucial role in modulating the growth and morphology of the cell. Harold’s team concludes that lower plants exhibit apical growth, a distinct mechanism where the cell wall expands solely at one end.
In summary, the realm of turgor pressure is replete with theoretical speculations that challenge traditional understandings. These theories, whether corroborated or refuted, contribute to the ever-evolving landscape of plant physiology.
Significance of Turgor Pressure
Turgor pressure, a fundamental concept in plant cellular biology, plays a pivotal role in maintaining the structural integrity and optimal functioning of plant cells. To elucidate its significance, let’s conceptualize two hypothetical cell systems: System A and System B.
System A: In this system, cells are densely packed, akin to bricks in a wall, with no intercellular spaces. Such cells have a high water concentration, resulting in elevated turgor pressure.
System B: Contrarily, cells in this system are loosely arranged with discernible spaces between them. These cells exhibit a flaccid nature, indicative of their low turgor pressure.
The disparity in turgor pressure between these two systems has profound implications for the overall health and appearance of the plant. Elevated turgor pressure, as seen in System A, ensures that the plant maintains its optimal shape and rigidity. This structural robustness is crucial for the plant to thrive in its environment and carry out essential physiological processes.
On the other hand, a decline in turgor pressure, as observed in System B, leads to the inward shrinking of cells. Given that plant cell walls possess limited flexibility, this shrinking results in the formation of intracellular spaces or voids. The manifestation of these voids at a macroscopic level is the wilting of the plant. Consequently, a plant with diminished turgor pressure appears droopy and unhealthy.
This correlation between turgor pressure and plant health underscores the importance of adequate water availability for plants. Water is not merely a medium for nutrient transport but is intrinsically linked to the plant’s structural integrity through turgor pressure. This pressure, observed not just in plants but also in fungi and certain bacteria, arises due to the osmotic movement of water across cell membranes.
In conclusion, turgor pressure is paramount for plants, ensuring their structural rigidity, optimal shape, and overall health. It serves as a barometer of the plant’s hydration status and, by extension, its well-being in its environment.
Functions of Turgor Pressure
Turgor pressure plays a crucial role in the physiology and functionality of plant cells. Here are the primary functions of turgor pressure:
- Structural Support: Turgor pressure provides rigidity to plant cells, helping maintain the structural integrity of the plant. This is especially important for non-woody plants that lack the rigid support of lignified tissues.
- Cell Expansion: During plant growth, turgor pressure aids in cell expansion. As water enters the cell, the increased turgor pressure pushes against the cell wall, leading to cell elongation and growth.
- Stomatal Regulation: Turgor pressure in the guard cells surrounding stomata regulates their opening and closing. This controls gas exchange (intake of carbon dioxide and release of oxygen) and transpiration in plants.
- Motility and Movement: Some plants exhibit rapid movements, such as the closing of the Venus flytrap or the folding of Mimosa leaves when touched. These movements are driven by changes in turgor pressure within specific cells.
- Prevention of Wilting: Adequate turgor pressure prevents plants from wilting. When plants are deprived of water, they lose turgor pressure, leading to drooping and wilting.
- Nutrient Uptake: Turgor pressure aids in the uptake of nutrients from the soil. As water moves into the root cells by osmosis, it brings along essential nutrients dissolved in it.
- Driving Force for Cell Division: In certain plant tissues, turgor pressure serves as a driving force for cell division, facilitating growth and development.
- Protection against Pathogens: Elevated turgor pressure can act as a barrier, preventing pathogens from penetrating and infecting plant cells.
In summary, turgor pressure is integral to various physiological processes in plants, from growth and movement to nutrient uptake and defense mechanisms. It ensures that plants remain upright, healthy, and responsive to their environment.
Quiz
What is turgor pressure primarily responsible for in plants?
a) Photosynthesis
b) Cellular respiration
c) Maintaining plant rigidity
d) Seed germination
In which organelle does the majority of water storage occur, contributing to turgor pressure in plant cells?
a) Chloroplast
b) Mitochondria
c) Vacuole
d) Nucleus
What happens to a plant cell when it loses turgor pressure?
a) It becomes turgid
b) It undergoes photosynthesis
c) It becomes flaccid
d) It divides rapidly
Which of the following is NOT a function of turgor pressure in plants?
a) Seed dispersal
b) Maintaining cell shape
c) DNA replication
d) Opening and closing of stomata
In which type of solution does a plant cell become turgid due to an increase in turgor pressure?
a) Isotonic
b) Hypertonic
c) Hypotonic
d) None of the above
Which component of the plant cell wall helps resist osmotic pressure and prevents the cell from bursting?
a) Lignin
b) Chitin
c) Cellulose
d) Starch
Turgor pressure is negative in which part of a transpiring plant?
a) Phloem
b) Xylem
c) Stomata
d) Guard cells
Which of the following organisms also regulate turgor pressure similar to plants?
a) Birds
b) Mammals
c) Bacteria
d) Fish
The movement of water molecules from an area of high water potential to an area of low water potential is known as:
a) Diffusion
b) Osmosis
c) Transpiration
d) Respiration
Which of the following is a result of low turgor pressure in plant cells?
a) The plant stands upright
b) The plant undergoes rapid growth
c) The plant appears wilted
d) The plant produces more seeds
FAQ
What is turgor pressure?
Turgor pressure is the force exerted by water inside the cell against the cell wall, providing structural support and rigidity to plants.
Why is turgor pressure important for plants?
Turgor pressure helps plants maintain their shape and rigidity. It also plays a crucial role in processes like the opening and closing of stomata and certain plant movements.
What happens when a plant cell loses turgor pressure?
When a plant cell loses turgor pressure, it becomes flaccid, leading to wilting in plants.
How is turgor pressure related to osmosis?
Turgor pressure arises due to the osmotic flow of water into the cell. When there’s a higher concentration of solutes inside the cell than outside, water moves into the cell by osmosis, increasing the turgor pressure.
What is the difference between turgid and flaccid cells?
A turgid cell is swollen and firm due to high turgor pressure, while a flaccid cell has lost its turgor pressure and is limp.
How do guard cells regulate turgor pressure to control stomatal opening?
Guard cells regulate the influx and efflux of potassium ions, which in turn affects the osmotic flow of water. When guard cells take in potassium ions, water follows by osmosis, increasing turgor pressure and causing the stomata to open. Conversely, when potassium ions are expelled, water exits the cells, reducing turgor pressure and causing the stomata to close.
How do non-plant organisms regulate turgor pressure?
Organisms like bacteria and fungi also regulate turgor pressure, primarily through modifications to their cell walls and by producing osmoprotectants or osmolytes to balance osmotic pressure.
What role does the cell wall play in turgor pressure?
The cell wall provides resistance to the expanding cell, preventing it from bursting due to excessive water influx. It helps maintain turgor pressure by providing rigidity and counteracting the osmotic pressure of water entering the cell.
How does turgor pressure influence plant movements?
Some plants exhibit movements in response to external stimuli, like the closing of the Mimosa pudica leaves when touched. These movements are driven by changes in turgor pressure within specialized cells.
Can turgor pressure be used for seed dispersal?
Yes, some plants, like the squirting cucumber (Ecballium elaterium), use turgor pressure for seed dispersal. The buildup of pressure inside the fruit causes it to burst, ejecting seeds at a distance.
References
- Biology Online. (Year of Publication). Turgor pressure. Retrieved from https://www.biologyonline.com/dictionary/turgor-pressure
- Biology Dictionary. (Year of Publication). Turgor pressure. Retrieved from https://biologydictionary.net/turgor-pressure/
- Hydrostatic Pressure. (2010). Retrieved from Engineeringtoolbox.com website: https://www.engineeringtoolbox.com/hydrostatic-pressure-water-d_1632.html
- CollegeDunia. (Year of Publication). Turgor pressure: Definition & explanation. Retrieved from https://collegedunia.com/exams/turgor-pressure-definition-explanation-biology-articleid-4517
- Science Facts. (Year of Publication). Turgor pressure. Retrieved from https://www.sciencefacts.net/turgor-pressure.html
- “pressure potential.” A Dictionary of Ecology. Retrieved from Encyclopedia.com: https://www.encyclopedia.com/science/dictionaries-thesauruses-pictures-and-press-releases/pressure-potential
- Kozlowski, T.T. (2012). Seed Biology: Importance, Development and Germination. 1. Academic Press. pp. 195–196.
- Keith, R. (2007). Handbook of Plant Science, Volume 1. John Wiley & Sons.
- Shimazaki, Y., Ookawa, T., & Hirasawa, T. (2005-09-01). “The Root Tip and Accelerating Region Suppress Elongation of the Decelerating Region without any Effects on Cell Turgor in Primary Roots of Maize under Water Stress”. Plant Physiology. 139 (1): 458–465.