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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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724 Chapter 13: Intracellular Membrane Traffic

hydrolase

intralumenal

vesicle

late endosome

endolysosome

Figure 13–39 A model for lysosome

maturation. Late endosomes fuse with

preexisting lysosomes (bottom arrow) or

preexisting endolysosomes (top arrow).

Endolysosomes eventually mature into

lysosomes as hydrolases complete the

digestion of their contents, which can include

intralumenal vesicles. Lysosomes also fuse

with phagosomes, as we discuss later.

digestion of

contents

lysosome

hydrolytically active, low-pH compartments

hydrolytic enzymes. It is especially hard to apply a narrower definition than this in

plant cells, as we discuss next.

Plant and Fungal Vacuoles Are Remarkably Versatile Lysosomes

Most plant and fungal cells (including yeasts) contain one or several very large,

MBoC6 m13.38/13.39

fluid-filled vesicles called vacuoles. They typically occupy more than 30% of the

cell volume, and as much as 90% in some cell types (Figure 13–40). Vacuoles are

related to animal cell lysosomes and contain a variety of hydrolytic enzymes, but

their functions are remarkably diverse. The plant vacuole can act as a storage

organelle for both nutrients and waste products, as a degradative compartment,

as an economical way of increasing cell size, and as a controller of turgor pressure

(the osmotic pressure that pushes outward on the cell wall and keeps the plant

from wilting) (Figure 13–41). The same cell may have different vacuoles with distinct

functions, such as digestion and storage.

The vacuole is important as a homeostatic device, enabling plant cells to withstand

wide variations in their environment. When the pH in the environment

drops, for example, the flux of H + into the cytosol is balanced, at least in part, by

an increased transport of H + into the vacuole, which tends to keep the pH in the

cytosol constant. Similarly, many plant cells maintain an almost constant turgor

pressure despite large changes in the tonicity of the fluid in their immediate environment.

They do so by changing the osmotic pressure of the cytosol and vacuole—in

part by the controlled breakdown and resynthesis of polymers such as

polyphosphate in the vacuole, and in part by altering the transport rates of sugars,

(A)

(B)

cell wall

vacuole

cytosol

chloroplasts

vacuole

tonoplast

10 µm

Figure 13–40 The plant cell vacuole.

(A) A confocal image of cells from an

Arabidopsis embryo that is expressing an

aquaporin—YFP (yellow fluorescent protein)

fusion protein in its tonoplast, or vacuole

membrane (green); the cell walls have been

false-colored orange. Each cell contains

several large vacuoles. (B) This electron

micrograph of cells in a young tobacco

leaf shows the cytosol as a thin layer,

containing chloroplasts, pressed against

the cell wall by the enormous vacuole.

(A, courtesy of C. Carroll and L. Frigerio,

based on S. Gattolin et al., Mol. Plant

4:180–189, 2011. With permission from

Oxford University Press; B, courtesy of

J. Burgess.)

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