Vacuole and Chloroplasts

To predict whether and in what direction water will move, we need to know the value of the water potential in the various compartments under consideration. At equilibrium, the water potential is the same in all communicating phases, such as those separated by membranes. For example, when water is in equilibrium across the tonoplast, the water potential is the same in the vacuole as it is in the cytosol. No force then drives water across this membrane, and thus no net flow of water occurs into or out of the vacuole. 

The tonoplast does not have an appreciable difference in hydrostatic pressure across it. A higher internal hydrostatic pressure would cause an otherwise slack (folded) tonoplast to be mechanically pushed outward. The observed lack of such motion indicates that AP is close to zero across a typical tonoplast. If the tonoplast were taut, AP would cause a stress in the membrane, analogous to the cell wall stresses discussed in Chapter 1 (Section 1.5C) namely, the stress would be rAPI2t for a spherical vacuole (see Eq. 1.15). However, the tensile strength of biological membranes is low—membranes can rupture when a stress of 0.2 to 1.0 MPa develops in them. For a tonoplast 7 nm thick with a maximum stress before rupturing of 0.5 MPa surrounding a spherical vacuole 14 pm in radius, the maximum hydrostatic pressure difference across the tonoplast is 

The central vacuole is a relatively simple aqueous phase that can act as a storage reservoir for metabolites or toxic products. For example, the nocturnal storage of organic acids, such as malic acid, takes place in the central vacuoles of Crassulacean acid metabolism plants (mentioned in Chapter 8, Section 8.5A), and certain secondary chemical products, such as phenolics, alkaloids, tannins, glucosides, and flavonoids (e.g., antho-cyanins), often accumulate in central vacuoles. Compared with the central vacuole, the cytoplasm is a more complex phase containing many colloids and membrane-bounded organelles. Because the central vacuole contains few colloidal or other interfaces, any matric pressure in it is 

In this chapter we will derive the Boyle-Van t Hoff relation using the chemical potential of water, and in Chapter 3 (Section 3.6B) we will extend the treatment to penetrating solutes by using irreversible thermodynamics. Although the Boyle-Van t Hoff expression will be used to interpret the osmotic responses only of chloroplasts, the equations that will be developed are general and can be applied equally well to mitochondria, whole cells, or other membrane-surrounded bodies. 

The Boyle-Van t Hoff relation applies to the equilibrium situation for which the water potential is the same on either side of the two membranes surrounding a chloroplast. When T1 equals T°, net water movement across the membranes ceases, and the volume of a chloroplast is constant. (The superscript i refers to the inside of the cell or organelle and the superscript o to the outside.) If we were to measure the chloroplast volume under such conditions, the external solution would generally be at atmospheric pressure (P° =0). By Equation 2.13a (T = P — H, when the gravitational term is ignored), the water potential in the external solution is then 

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Topical eukaryotic cells (Fig. 1-7) are much larger than prokaryotic cells—commonly 5 to 100 pm in diameter, with cell volumes a thousand to a million times larger than those of bacteria. The distinguishing characteristics of eukaryotes are the nucleus and a variety of membrane-bounded organelles with specific functions mitochondria, endoplasmic reticulum, Golgi complexes, and lysosomes. Plant cells also contain vacuoles and chloroplasts (Fig. 1-7). Also present in the cytoplasm of many cells are granules or droplets containing stored nutrients such as starch and fat. 

Most eukaryotic cells contain many mitochondria, which occupy up to 25 percent of the volume of the cytoplasm. These complex organelles, the main sites of ATP production during aerobic metabolism, are generally exceeded in size only by the nucleus, vacuoles, and chloroplasts. 

An H+ electrochemical gradient (ApH+) provides the energy required for active transport of all classical neurotransmitters into synaptic vesicles. The Mg2+-dependent vacuolar-type H+-ATPase (V-ATPase) that produces this gradient resides on internal membranes of the secretory pathway, in particular endosomes and lysosomes (vacuole in yeast) as well as secretory vesicles (Figure 3). In terms of both structure and function, this pump resembles the F-type ATPases of bacteria, mitochondria and chloroplasts, and differs from the P-type ATPases expressed at the plasma membrane of mammalian cells (e.g., the Na+/K+-, gastric H+/K+-and muscle Ca2+-ATPases) (Forgac, 1989 Nelson, 1992). The vacuolar and F0F1... 

A eukaryotic cell is surrounded by a plasma membrane, has a membrane-bound nucleus and contains a number of other distinct subcellular organelles (Fig. 1). These organelles are membrane-bounded structures, each having a unique role and each containing a specific complement of proteins and other molecules. Animal and plant cells have the same basic structure, although some organelles and structures are found in one and not the other (e.g. chloroplasts, vacuoles and cell wall in plant cells, lysosomes in animal cells). 

According to this hypothesis (Margulis, 1993), the eukaryotic cell is a result of symbiosis of different prokaryotic cells, where mitochondria originated from eubacteria, and chloroplasts - from cyanobacteria, and vacuoles - from archae. 

The rate of photosynthesis does not depend on the amount of a single component (e.g., the activity of a particular enzyme). There is a wide range of possible regulatory factors, proven to exist in vitro, but the importance of which in vivo has still to be determined. In particular, there is a multitude of factors affecting the activity of the enzymes involved, with pH, ions, coenzymes, and metabolite effectors modulating the activity of every enzyme studied thus far. Compartmentation is the other key factor. The role of metabolite transport in the cell, particularly between chloroplast and cytosol, but also to and from mitochondria, vacuole, and other organelles, is now considered to be fundamental to the regulation of photosynthesis. In this chapter, we look at the factors considered to be of major importance... 

The structures of F class and V class ion pumps are sIm liar to one another but unrelated to and more complicated than P-class pumps. F- and V-class pumps contain several different transmembrane and cytosolic subunits. All known V and F pumps transport only protons. In a process that does not Involve a phosphoprotein Intermediate. V-class pumps generally function to maintain the low pH of plant vacuoles and of lysosomes and other acidic vesicles In animal cells by pumping protons from the cytosolic to the exoplasmic face of the membrane against a proton electrochemical gradient. F-class pumps are found In bacterial plasma membranes and In mitochondria and chloroplasts. In contrast to V pumps, they generally function to power the synthesis of ATP from ADP and Pj by movement of protons from the exoplasmic to the cytosolic face of the membrane down the proton electrochemical gradient. Because of their Importance In ATP synthesis in chloroplasts and mitochondria, F-class proton pumps, commonly called ATP synthases, are treated separately In Chapter 8. 

Unique subcellular compartmentation is also present in quinolizidine alkaloid biosynthesis, which occurs in the mesophyll chloroplasts of some legumes (177). HMT/HLT activity is localized to the mitochondrial matrix, but not to chloroplasts where de novo quinolizidine alkaloid biosynthesis is thought to occur (176). ECT is present in an organelle distinct from the mitochondria and chloroplasts, but has not been unambiguously localized. Although the quinolizidine nucleus appears to be synthesized in the chloroplast, subsequent modifications can occur only after alkaloid intermediates are transported to the cytosol and mitochondria. Quinolizidine alkaloids appear to accumulate in vacuoles of epidermal cells where their defensive properties would be most effective. 

Several products which are known to be synthesized in plastids are eventually released into extraplastid compartments. Flavonoids or chlorogenic acid may be formed in chloroplasts and transported from there to their storage sites in vacuoles and the periplasmic space. In certain cases, the release of secondary products from chloroplasts seem to be strictly regulated such as the flow of the chloroplast-synthesized gibberellic acids to the cytoplasm, which is controlled by phytochrome. 

Those succulents which unify the sites of malic acid synthesis, storage (large vacuoles), and conversion (chloroplasts) all within the same cells, can be expected to have CAM (see Fig. 2.3). In contrast, those succulents where the potential malic acid stores (water cells) are spatially separated from the sites of photosynthesis will not or only very weakly perform CAM, probably because of the complexity of transport of malic acid. 

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