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Chapter 1. The Scene of Action

make up the pith or pulp in the center of stems and roots, where they serve as food storage cells. The collenchyma, present in herbs, is composed of elongated supporting cells and the sclerenchyma of woody plants is made up of supporting cells with hard lignified cell walls and a low water content. This tissue includes fiber cells, which may be extremely long; e.g., pine stems contain fiber cells of 40 m diameter and 4 mm long. Two complex tissues, the xylem and phloem, provide the conducting network or circulatory system of plants. In the xylem or woody tissue, most of the cells are dead and the thick-walled tubes (tracheids) serve to transport water and dissolved minerals from the roots to the stems and leaves. The phloem cells provide the principal means of downward conduction of foods from the leaves. Phloem cells are joined end to end by sieve plates, so-called because they are perforated by numerous minute pores through which cytoplasm of adjoining sieve cells appears to be connected by strands 5 9 m in diameter.154 Mature sieve cells have no nuclei, but each sieve cell is paired with a nucleated companion cell. Epidermal tissue of plants consists of flat cells, usually containing no chloroplasts, with a thick outer wall covered by a heavy waxy cuticle about 2 m thick. Only a few specialized cells are found in the epidermis. Among them are the paired guard cells that surround the small openings known as stomata on the undersurfaces of leaves and control transpiration of water. Specialized cells in the root epidermis form root hairs, long extensions (~1 mm) of diameter 517 m. Each hair is a single cell with the nucleus located near the tip. Figure 1-16 shows a section from a stem of a typical angiosperm. Note the thin cambium layer between the phloem and the xylem. Its cells continuously

undergo differentiation to form new layers of xylem increasing the woody part of the stem. New phloem cells are also formed, and as the stem expands all of the tissues external to the cambium are renewed and the older cells are converted into bark. Plant seeds consist of three distinct portions. The embryo develops from a zygote formed by fusion of a sperm nucleus originating from the pollen and an egg cell. The fertilized egg is surrounded in the gymnosperms by a nutritive layer or endosperm which is haploid and is derived from the same gametophyte tissue that produced the egg. In angiosperms two sperm nuclei form; one of these fertilizes the egg, while the other fuses with two haploid polar nuclei derived from the female gametophyte. (The polar nuclei are formed by the same mitotic divisions that formed the egg.) From this develops a 3n triploid endosperm. G. The Chemical Composition of Cells Water is the major component of living cells, but the amount varies greatly. Thus, the pig embryo is 97% water; at birth a new-born pig is only 89% water. A lean 45-kg pig may contain 67% water but a very fat 135-kg animal only 40% water. Similar variations are encountered with other constitutents. The water content of a tissue is often determined by thoroughly drying a weighed sample of tissue at low temperature in vacuum and then weighing it a second time. The solid material can then be extracted with a solvent that will dissolve out the fatty compounds. These are referred to collectively as lipids. After evaporation of the solvent the lipid residue may be weighed. By this procedure a young leafy vegetable might be found to contain 2 5% lipid on a dry weight

Ray

Ray
Sieve tube

Ray

Phloem Cambium Xylem

Fiber Sieve plate

Pith

Vessel Fiber

Figure 1-16 Section of the stem of an angiosperm. Enlarged sections showing tubes of the phloem (left) and xylem (right). From S. Biddulph and O. Biddulph.155 Drawn by Bunji Tagawa.

G. The Chemical Composition of Cells

31

basis. Even very lean meats contain 10 30% lipid. The residue remaining after removal of the lipid consists predominately of three groups of compounds: proteins, nucleic acids, and carbohydrates. Most of the nitrogen present in tissues is found in the proteins and the protein content is sometimes estimated by determining the percentage of nitrogen and multiplying by 6.25. In a young green plant, 20 30% of the dry matter may be protein, while in very lean meat it may reach 50 70%.

TABLE 1-4 Approximate Composition of Metabolically Active Cells and Tissuesa

Green plant (spinach, Spinacia oleracea)c 93 2.3 Rat liverd (%) 69 21

Component H2O Protein Amino acids DNA RNA Nucleotides Carbohydrates Cellulose Glycogen Lipids Phospholipids Neutral lipids Sterols Other small molecules Inorganic ions K+

E. colib (%) 70 15 0.4 1 6 0.4 3

0.2 1.0

3.2 0.6 0.3

3.8 6 3.1 1.6 0.3

0.2 1 1.5 0.4

Equivalents per liter in rat liver Amino acid residues Nucleotide units Glycogen (glucose units) K+
a b

2.1 0.03 0.22 0.1

c d

Data were not readily available for spaces left blank From J. D. Watson (1976) Molecular Biology of the Gene, 3rd ed., p. 69, Benjamin, New York The amounts of amino acids, nucleotides, carbohydrates, and lipids include precursors present in the cell. From B. T. Burton (1976) Human Nutrition, 3rd ed., McGraw-Hill, New York (p. 505) From C. Long, ed., (1961) Biochemists Handbook, pp. 677 679, Van Nostrand-Reinhold, Princeton, New Jersey

A dried tissue sample may be burned at a high temperature to an ash, which commonly amounts to 310% and is higher in specialized tissues such as bone. It is a measure of the inorganic constituents of tissues. The carbohydrate content can be estimated by the difference of the sum of lipid, protein, and ash from 100%. It amounts to 50 60% in young green plants and only 2 10% in typical animal tissues. In exceptional cases the carbohydrate content of animal tissues may be higher; the glycogen content of oysters is 28%. The amount of nucleic acid in tissues varies from 0.1% in yeast and 0.5 1% in muscle and in bacteria to 1540% in thymus gland and sperm cells. In these latter materials of high nucleic acid content it is clear that multiplication of % N by 6.25 is not a valid measure of protein content. For diploid cells of the body the DNA content per cell is nearly constant. Table 1-4 compares the composition of a bacterium, of a green plant, and of an active animal tissue (rat liver). Although the solid matter of cells consists principally of C, H, O, N, S, and P, many other chemical elements are also present. Among the cations, Na+, K+, Ca2+, and Mg2+ are found in relatively large amounts. Thus, the body of a 70 kg person contains 1050 g Ca (mostly in the bones), 245 g K, 105 g Na, and 35 g Mg. Iron (3 g), zinc (2.3 g), and rubidium (1.2 g) are the next most abundant. Of these iron and zinc are essential to life but rubidium is probably not. It is evidently taken up by the body together with potassium. The other metallic elements in the human body amount to less than 1 g each, but at least seven of them play essential roles. They include copper (100 mg), manganese (20 mg), and cobalt (~5 mg). Others, such as chromium (<6 mg), tin, and vanadium, have only recently been shown essential for higher animals.156,157 Nickel, lead, and others may perhaps be needed. Nonmetallic elements predominating in the ash are phosphorus (700 g in the human body), sulfur (175 g), and chlorine (105 g). Not only are these three elements essential to all living cells but also selenium, fluorine, silicon (Box 4-B), iodine, and boron are needed by higher animals and boron by plants (Fig. 1-17). Iodine deficiency may affect one billion human beings and may cause 20 million cases per year of cretinism, or less severe brain damage.158 What is the likelihood that other elements will be found essential? Consider a human red blood cell, an object of volume ~80 m3 and containing about 3 x 108 protein molecules (mostly hemoglobin). About 7 x 105 atoms of the trace metal copper and 105 atoms of the nutritionally essential tin are present in a single red cell. Also present are 2 x 104 atoms of silver, a toxic metal. Its concentration, over 10-7 M, is sufficient that it could have an essential catalytic function. However, we know of none and it may simply have gotten into our bodies from handling money, jewelry, and other