Cell membranes and ion transport

chapter 1
structure and properties of cell membranes
all living cells are delimited from their environment by a surface membrane, the plasma membrane or
plasmalemma, while eukaryotic plant and animal cells are further compartmentalized intracellularly into
various membrane bound organelles. Perhaps the most fundamental property of these membranes is
their ability to act as selective permeability barriers , controlling the amount and nature of the
substances that pass across them. The development of such a boundary was essential to the
development of primitive cells , which presumably arose by the formation of a membrane around a few
macromolecules possessing catalytic properties, thus allowing other smaller molecules and ions to
interact. Such a membrane would need to possess a certain selectivity allowing the cell to develop
characteristic properties; it would permit the selection and retention of essential molecules that became
depleted and a removal of waste products of metabolism. Such transport systems must have appeared
quite early in evolution, perhaps before the appearance of metabolic pathways see Holden, 1968 ;
Pardee and Palmer , 1973). Today, organelles and pleuropneumonia-like organisms which lives
surrounded by cytoplasm possess membranes and transport systems but lack a complete enzymic
machinery, an observation consistent with the above idea. If this is the case, then presumably
membranes evolved even earlier. In addition, such a cell would need the ability to respond to changes in
the external medium. For example, the cell volume must remain fairly constant, since large changes
would affect the concentration of molecules within the cell and so the interactions between them. This
could be agent to counteract the changes occurring in the environment. Today , many animal cells
regulate their osmotic environment by regulating the internal concentration of sodium ions.
the permeability of cells membranes to different substances varies widely. Gases move across
membranes quite easily and small molecules pass through more easily than larger molecules with
similar chemical properties. Usually the grater the lipid solubility of a substance, the greater its
permeability through the membrane, while cells are usually less permeable to electrolytes than non-
electrolytes. Substances may diffuse through membranes passively or may require the expenditure of
energy to achive this movement. The selectivity of membranes prevents the random diffusion of
substances and allow the cell to determine and control is internal environment while providing a
number of different micro- environment within the subcellular organelles. Membranes allow the cell or
cell prganelle to be isolated from its surroundings and so allow the highly ordered biochemical
processes of life to go on relatively un affected by changes in the environment. At the same time,
membranes allow a constant exchanges of substances with the outside medium, including the uptake of
oxygen and essential nutrients and the exclusion of un wanted and harmful materials. The control of ion
movement is clearly of fundamental importance in this homeostatic process since it will directly affect
such factors are cellular Ph and various osmotic and metabolic functions. Animals cells, in particular
show remarkably consistent electrolyte contents, and both plant and animalcellsmaintain concentration
of individual ions which are very different from those in the extracellular environment. The most
oimportant membrane in this respect is theplasma membrane, which is the primary barrier to the
movement of ions into and out of cells, . in vacuolated plant cells, the tonoplast or vacuator membrane
provides a second important provides a second important barrier, since the vacuole is a major site of ion
accumulation . in addition, the ion content of cell organelles may show marked differences from that of
other parts of the cell and environment . thus, when we consider the total ionic relations of a cell tissue,
it must be remembered that more thanone membrane barrier is involved , thewhole providing a
complex system of selectively permeable barriers. However , although we will be largely concerned here
with the movement of ions, this is only one of a wide variety of activities associated with membrane
structures. Membranes areinvolved in functions such as cell-cell interactions, contractile processes,
excitation in nerves, ATP generation , and energy conversion in photosynthesis. Many of these processes
are in turn closely related to the movement of ions across membranes.
Composition of memberanes
our knowledge of membrane structure and composition has increased enormously in the last 20 years
with the development of electron microscopy and modern biochemical techniques. However, it should
be remembered that much was learnt of the nature of membranes long before remembered that much
was learnt of the nature of membranes long before they could readily be distinguished by electron
microscopy. The presence of a cell membrane or plasma membrane was inferred by the early light
microscopists from observations of the swelling and contraction of cells, the uptake of dyes, and the loss
of contents when the cell surface is torn. Electron microscopy confirmed the universal existence of this
membrane. However , even today, the selective role of the plasma membrane, particularly in relation to
ions, is doubted by minority of authors who claim that selectivity is a property of the cytoplasm. This
view will be discussed more fully later.
One of the earliest predictions as to the nature of this membrane came from permeability
studies of plant cells conducted by overton towards the end of the nineteenth century. He
demonstrated that the rate of penetration of non- electrolytes was correlated with their relative
solubility in fats and so concluded that plasma membrane was essentially lipoidal in nature. These
observations werw strengthened by the classic studies of colander some 40 years later on the
permeability of giant algal cells. An approximately linear relationship was found between the oil
solubility of various solutes and a parameter PM
1,5
which combines the permeability and molecular size
of the solute. This relationship is discussed more fully in chapter 3. Further evidence for the lipoidal
nature of cel membranes came from studies of their electrical properties, which were similar to these of
lipid layers. However this was not the whole story. In the 1935 Danielli and Davson reported that the
surface tension of cells was much lower than the high values obtained for neutral lipid- water interfaces,
suggesting that protein was present at the membrane surface. The amphoteric nature of cell surfaces,
which Is also a characteristic property of protein molecules, and the fact that proteases could destroy
cell membranes lent further support to the concept that protein was an essential component of cell
membranes. Subsequent analysis of certain membranes isolated free from other cellular materials has
shown that they consist largely of lipid and protein, together with some carbohydrate, thus confirming
the prediction based on physiological and physical measurements. The carbohydrate is in the from of
sugar residues attached to protein or lipid to form glycoproteins and glycolipids.
The chemical composition of membranes is not easy to determine since