TIME RELATIONS OF GROWTH : III. GROWTH CONSTANTS DURING THE SELF-ACCELERATING PHASE OF GROWTH.

Growth curves consist, in all cases, of two major segments. The first major segment is, in the case of higher animals and plants, made up in turn of several (probably five) shorter segments during each of which growth takes place at a constant percentage rate. The transitions between the successive stages are abrupt, the abruptness being of the order of metamorphosis in cold blooded animals. It has been made clear in the first paper of this series that the time rate of growth following the major inflection declines at a constant percentage rate. The junction between the two major segments occurs at puberty in animals and flowering in plants. The two major segments are not symmetrical about the major inflection. The slope of the segment following the inflection is always less than the slope of the segment preceding the inflection. The major inflection does not occur in the center of the growth curve. The instantaneous rate of growth at the beginning of growth is of the order of 100–200 per cent per day (i.e. the body weight is doubled in from 7 to 17 hours). It may be mentioned that 2 months after conception the rate of growth in man is only 8 per cent per day. This is contrary to all the published statements. Thus, Minot concluded that growth begins at 1000 per cent per day; Jackson concluded that in man, growth during the 1st month takes place at 57.5 million per cent per month; during the 2nd month 990 per cent per month; during the 3rd month 390 per cent per month (8 per cent per day is only 240 per cent per month). The reason for the discrepancy between the values derived, by the method adopted by the writer, and the values given in the literature is explained by Fig. 1.     

THE RATE OF GROWTH OF THE DAIRY COW : V. EXTRAUTERINE GROWTH IN LINEAR DIMENSIONS.

Barring fluctuations due to the cyclic phenomena, the extrauterine course of growth in linear dimensions and in weight of the dairy cow follows an exponential law having the same form as the law representing the course of monomolecular change in chemistry. This suggests the interpretation that the general course of growth is limited by a monomolecular chemical process, and that the cyclic phenomena are due to subsidiary processes in the fundamentally exponential course of growth. The fact that growth follows or tends to follow an exponential course may be stated more simply as follows: if the unit of time is taken sufficiently large so that fluctuations due to the cyclic phenomena are balanced or eliminated, then the amount of growth made during the given unit of time at any age tends to be a constant percentage of the growth made during the preceding unit of time. Thus, the growth in height at withers made during any year is about 34 per cent of the growth made during the preceding year. Similarly the growth in weight made during any year is about 56 per cent of the growth in weight made during the preceding year. This is in accordance with expectations if it is assumed that each animal begins life with a definite endowment of limiting substance necessary for the process of growth, and that this endowment is used up at a constant rate (or percentage) of itself.     

THE RATE OF GROWTH OF THE DAIRY COW : II. GROWTH IN WEIGHT AFTER THE AGE OF TWO YEARS.

An extensive amount of data is presented on the growth in weight of the dairy cow from 2 to 17 years of age, covering practically the entire duration of life. The data show that after the age of 2 years the rate of growth declines in a non-cyclic manner. The course of decline in growth follows the course of decline of a monomolecular chemical reaction; that is, the percentage decline in growth with age is constant.     

THE RATE OF GROWTH OF THE DAIRY COW : EXTRAUTERINE GROWTH IN WEIGHT.

The growth period of the Jersey and Holstein cows is made up of at least three cycles, two extrauterine cycles with maxima at about 5 and 20 months of age, and one intrauterine cycle, the maximum of which has not yet been determined. The equation of an autocatalytic monomolecular reaction was found to give very good results when applied to the cycle having its maximum at about 5 months of age. The values obtained from this equation when applied to the cycle having the maximum at about 20 months of age were higher than the observed values probably due to the retarding effect of pregnancy and lactation on growth.     

THE NATURE OF THE GROWTH RATE

1. The growth rate of organisms may be considered as a chemical reaction which gives the mature organism as its end-product. The organism grows at a definite rate which is, at any moment, proportional to the amount of growth yet to be made. 2. Shoots of young pear trees measured at weekly intervals during the growing season showed a rate similar to that of an autocatalytic reaction. 3. Young walnut trees showed distinct cycles of growth in a single season, but the growth in each cycle proceeded at a rate corresponding to an autocatalytic reaction. 4. The growth rate follows a definite, quantitative course though judged by different criteria. Data are presented for maize in which green weight, dry weight, and height of the plant are used. Data for cattle show that either weight or height of the animal may be used as a criterion.     

STUDIES ON THE GROWTH HORMONE OF PLANTS : VI. THE DISTRIBUTION OF THE GROWTH SUBSTANCE IN PLANT TISSUES

1. It is shown that when plant tissues are ground with water the growth substance contained therein is inactivated by the oxidizing enzymes. 2. A simple method of extraction is described which enables the quantitative determination of growth substance in such tissues. 3. The amount and distribution of growth substance in the Avena coleoptile is determined by this method, and it is shown that while the substance does not diffuse out from the lower parts of the coleoptile, it is nevertheless present in considerable amounts, the concentration decreasing steadily with the distance from the tip. 4. Growth substance is also present in considerable amounts in Avena roots, and here also its concentration decreases steadily with distance from the tip. 5. The amount of growth substance diffusing out of root tips into dextrose agar, even during long periods of time, is not greater than the amount obtainable by direct extraction. Actual production in the root tip therefore either does not take place at all, or else takes place under quite different conditions from the production in the tip of the coleoptile.     

THE ACTION OF THE PLANT GROWTH HORMONE

1. Sections of Avena coleoptiles are found to show a considerable elongation when suspended in solutions of growth substance. 2. This elongation does not take place in the absence of O₂ and is inhibited by KCN and phenylurethane. 3. The rate of respiration of sections of coleoptiles is increased by the addition of growth substance in concentrations which cause growth. High concentrations of growth substance inhibit growth and also respiration. 4. The increase in respiration is inhibited by KCN and phenylurethane in the concentrations which inhibit normal respiration. These concentrations are the same as those which inhibit growth. 5. From 2, 3, and 4, it seems possible that the increase in respiration caused by growth substance may be an essential part of its action in growth.     

THE RATE OF GROWTH OF THE DAIRY COW : III. THE RELATION BETWEEN GROWTH IN WEIGHT AND INCREASE OF MILK SECRETION WITH AGE.

It is shown that from 2 years, the age when milk secretion usually begins, to 9 years, the age of maximum body weight, the increase of milk secretion with age follows the course of growth in body weight— both can be accurately represented by the equation of a monomolecular chemical reaction having a velocity constant of approximately the same numerical value. While increase in milk secretion and increase in body weight with age follow the same course, it is shown that increasing body weight contributes only about 20 per cent to increasing milk secretion with age. The fact that milk secretion and body weight follow the same course, even though they are largely independent of each other indicates that increase in body weight is a good measure of growth of the dairy cow; this fact also shows that the increase of milk secretion with age may be used as a measure of growth. The fact that milk secretion, like body weight, follows the course of a chemical reaction, adds further support to the theory that growth is limited by a chemical reaction.     

THE LIGHT GROWTH RESPONSE AND THE GROWTH SYSTEM OF PHYCOMYCES

1. The Roscoe-Bunsen law holds for the light growth response of Phycomyces if the time component of stimulation is short. With exposures longer than a few seconds, the reaction time to light is determined by the intensity and not by the energy of the flash. 2. The possible nature of the very long latency in the response to light is considered in terms of the structure of the cell and its mechanism of growth. It is suggested that during the latency some substance produced by light in the protoplasm is transported centrifugally to the cell wall or outermost layer of protoplasm. 3. The total elongation occurring over a period of 1 to 2 hours is independent of flashes of light or temporary darkening. Light acts by facilitating some change already under way in the growth system, and during the principal phase of elongation is not a necessary or limiting factor for growth. 4. Judged by the reaction time, the original sensitivity is restored in the light system following exposure to light in about one-third the time required for equilibrium to be reattained in the growth system.     

不同营养方式对小球藻FACHB 484生长的影响及其非自养生长机理研究

系统研究了小球藻FACHB 484在含有葡萄糖的不同营养方式下的生长情况,并通过抑制试验探讨葡萄糖在小球藻FACHB 484光异养和兼养生长条件下所起的作用以及小球藻FACHB 484是否存在氧化呼吸系统的关键酶类。结果表明:小球藻FACHB 484可利用葡萄糖进行化能异养、光激活异养、光异养及兼养生长,其生长速率大小为:兼养光异养光激活异养化能异养光合自养。兼养培养的最大生物量和比生长速率分别是自养培养的8.6和3.4倍,其比生长速率接近于光合自养和光异养培养下的比生长速率之和。葡萄糖主要作为小球藻FACHB 484兼养和光异养培养的碳源,而能量主要源自光。小球藻FACHB 484存在氧化呼吸链代谢途径,其细胞中有琥珀酸脱氢酶和细胞色素氧化酶。       

THE PRENATAL GROWTH OF THE MOUSE

1. The general course of prenatal growth in the mouse, the guinea pig, and the chick can be expressed by straight line relations between the logarithms of the weight and age only when age is counted from the beginning of the embryo proper. 2. This is interpreted as showing that the manner of growth before the beginning of the embryo proper is essentially different from that after this time. 3. The velocity constants for the animals mentioned are similar; the major differences in their curves depend on the amount of tissue involved in the first organization of the embryo proper and in the length of prenatal life. 4. Growth of different animals may be compared more accurately if, instead of either birth age or conception age, embryo age is used.     

GROWTH AND ORGANIZED DEVELOPMENT OF CULTURED CELLS IV. THE BEHAVIOR OF THE NUCLEUS

Mitra , J., Marion O. Mapes , and F. C. Steward . (Cornell U., Ithaca, New York.) Growth and organized development of cultured cells. IV. The behavior of the nucleus. Amer. Jour. Bot. 47(5) : 357—368. Illus. 1960.–The nuclei and the chromosomes of carrot cells have been examined at various stages throughout the following sequence: (1) growth of a tissue culture from a preformed explant of secondary phloem from the carrot root; (2) growth and multiplication of carrot cells freely suspended in a liquid medium; (3) growth and re-formation of organs (roots) and whole plants (including flowers) from cells in the freely suspended state. The cells of the carrot are normally diploid (2n = 18), the cells which develop in the explant are also diploid, and the cells of the re-formed organs, and even the flowers developed upon plants grown from cells, are also normal and diploid; normal meioses also occur. Nevertheless, the wide range in growth and form of the freely suspended cells is accompanied by a rich diversity of cytological conditions; these include tetraploid and highly polyploid nuclei which divide, haploidy and such chromosomal aberrations as di- and even tri-centric bridges. Two division figures showing chromosome numbers at different levels of ploidy were seen within the confines of one large cell, and, in another, 2 adjacent division figures were observed with chromosome numbers lower than diploid. Small thick-walled, densely protoplasmic cells divide to form bi- and tetra-nucleate conditions, and in a giant cell a highly multinucleate condition has been seen. Despite this, however, all the regenerated roots and plants yet examined are normally diploid. The implications of these events are discussed.     

THE GROWTH AND NUTRITION OF THE AFRICAN ELEPHANT

In order to gain information on the growth and nutrition of two populations of elephants in Murchison Falls National Park, Uganda, their urinary excretions of hydroxyproline peptides (HP) were compared. Animals from the “south bank” population had a consistently higher level of excretion than corresponding animals from the “north bank” population. This indicated a faster rate of growth and a superior nutritional status in “south bank” animals at the time of sampling. The HP output from both populations was found to run parallel to the seasonal variation in rainfall and vegetation growth. Sampling from each population was carried out at a different time of the year and evidence is presented to show that the differences in the rate of growth are a direct result of this. It is concluded that any primary differences between the populations are small enough to be obliterated by seasonal changes. The effect of the seasons on the growth of the elephant is therefore examined in more detail, and a direct influence on collagen metabolism is discussed.     

GROWTH OF THE EMBRYO OF GINKGO BILOBA UNDER EXPERIMENTAL CONDITIONS. III. GROWTH RATES OF ROOT AND SHOOT UPON MEDIA ABSORBED THROUGH THE COTYLEDONS

Ball , Ernest . (North Carolina State Coll., Raleigh.) Growth of the embryo of Ginkgo biloba under experimental conditions. III. Growth rates of root and shoot upon media absorbed through the cotyledons. Amer. Jour. Bot. 46(2) : 130-139. Illus. 1959.—Mature embryos of Ginkgo biloba were grown by inserting the cotyledons into agar medium containing one of the naturally-occurring sugars of the seed. These sugars were utilized at 1, 2, 4, 8 or 16% (w/v). Root growth was best on the sugar-mineral-water media and occurred at its maximum in media containing 4% sugar (ca. 0.25 M). The sugars may be listed as follows in order of decreasing effectiveness in root growth: glucose, sucrose, levulose, raffinose, galactose. Since very different growth rates of roots occurred on media of the same osmotic values when the latter were determined by the various sugars, it is suggested that the root utilized the entire sugar molecule as a unit whether it was mono-, di-, or trisaccharide. Such media were not effective in supporting shoot growth. Outstanding growth of shoots was obtained when either glutamine or coconut milk was placed in the culture medium. It is concluded that root growth can occur at a substantial rate by utilizing sugar and manufacturing organic nitrogen from nitrates. Shoot growth, in contrast, appears to require sources of organic nitrogen such as may be furnished by glutamine or coconut milk. These phenomena are thought to constitute fundamental differences between the metabolism of root and shoot of this embryo. Mannitol caused severe inhibition of growth of both root and shoot. It is probably toxic to this embryo. Indoleacetic acid caused severe inhibitions of growth of both root and shoot at all except the lowest concentrations utilized.     

THE PHOTOINHIBITION OF GROWTH IN ETIOLATED STEM SEGMENTS. II. GROWTH CAUSED BY COBALT IN PISUM

Bertsch, Walter F. (Yale U., New Haven, Conn.) The photoinhibition of growth in etiolated stem segments. II. Growth caused by cobalt in Pisum. Amer. Jour. Bot. 50(3): 213–219. Illus. 1963.—Etiolated pea stem sections were utilized to study the effects of various metal ions on the red, far-red photoinhibition of stem growth. Excision of immature tissue results in loss of photosensitivity. All growth caused by cobalt, either in the presence or absence of sugar, was inhibited by brief exposure of the excised tissue to red light. Since growth caused by sugar is also light-sensitive, it may be said that either cobalt or sugar may induce photosensitivity in this immature tissue. Other divalent metal ions were tested, but only nickel promoted growth. Although cobalt and nickel showed the same optimal concentration (ca. 2 × 10⁻⁵ m), growth caused by nickel was not photosensitive, suggesting that cobalt and nickel promoted growth through separate mechanisms. Since growth caused by indole-3-acetic acid or gibberellic acid is also insensitive to light in this tissue, a photosensitive sugar-cobalt growth system has been experimentally separated from other growth-controlling systems. The photoinhibition of a growth system which utilizes sugars and is promoted by cobalt and the relation of this system to certain de-etiolation phenomena are discussed.     

THE EFFECT OF AERATION ON THE GROWTH OF SPARTINA ALTERNIFLORA LOISEL.

A greenhouse experiment was designed to investigate the correlations between waterlogging and aeration, and associated changes in pH, redox potentials and sulfide concentrations, on the growth of Spartina alterniflora Loisel. Elemental concentrations of the aerial and root material were determined and used for correlations with growth response. Redox potentials adjusted to pH 7 (Eh 7) ranged from −184 mv to 5 mv and were highly correlated (r) with aerial and root dry weight biomass (.97 and .97, respectively) and plant height (1.0). The range of soil pH at the conclusion of the study was 6.07 to 6.74 and was negatively correlated with aerial and root dry weight biomass. Sulfide concentrations ranged from 10–² to 10–⁷ m and vorrelations with aerial and root dry weights and height were −.85, −.85 and −.87, respectively. High negative correlations were found between sodium and sulfur concentrations and S. alterniflora growth. Positive correlations between potassium, phosphorus, manganese, zinc, copper, iron and growth response were also observed. Correlations of elemental concentrations of the plants with redox potentials and/or pH suggest that these two physical variables may be responsible in part for the regulation of 5. alterniflora growth in nature by regulating availability of nutritional elements.     

THE RATE OF GROWTH OF THE DOMESTIC FOWL

This paper points out the fact that the growth period of the domestic fowl is analogous to that of the mammal, being composed of three, or perhaps four, cycles; two of these cycles are postembryonic with maxima at about 8 and 18 weeks varying somewhat with the breed and two or at least one, are embryonic with maxima at 11 to 12 and 15 to 16 days of age. Hatching occurs during the first part of the second or third cycle resembling in this respect the guinea pig rather than the mouse. The velocity curves of each of these cycles are similar to and can be represented by the equation of an autocatalytic monomolecular reaction.     

THE GROWTH OF BACTERIOPHAGE

1. An anti-Escherichia coli phage has been isolated and its behavior studied. 2. A plaque counting method for this phage is described, and shown to give a number of plaques which is proportional to the phage concentration. The number of plaques is shown to be independent of agar concentration, temperature of plate incubation, and concentration of the suspension of plating bacteria. 3. The efficiency of plating, i.e. the probability of plaque formation by a phage particle, depends somewhat on the culture of bacteria used for plating, and averages around 0.4. 4. Methods are described to avoid the inactivation of phage by substances in the fresh lysates. 5. The growth of phage can be divided into three periods: adsorption of the phage on the bacterium, growth upon or within the bacterium (latent period), and the release of the phage (burst). 6. The rate of adsorption of phage was found to be proportional to the concentration of phage and to the concentration of bacteria. The rate constant k_a is 1.2 x 10^–9 cm.⁸/min. at 15°C. and 1.9 x 10^–9 cm.⁸/min. at 25°. 7. The average latent period varies with the temperature in the same way as the division period of the bacteria. 8. The latent period before a burst of individual infected bacteria varies under constant conditions between a minimal value and about twice this value. 9. The average latent period and the average burst size are neither increased nor decreased by a fourfold infection of the bacteria with phage. 10. The average burst size is independent of the temperature, and is about 60 phage particles per bacterium. 11. The individual bursts vary in size from a few particles to about 200. The same variability is found when the early bursts are measured separately, and when all the bursts are measured at a late time.     

GROWTH AND ORGANIZED DEVELOPMENT OF CULTURED CELLS. V. THE GROWTH OF COLONIES FROM FREE CELLS ON NUTRIENT AGAR

When angiosperm cells are cultured in a liquid medium they may grow in the form of free, single cells and form small to large groups of cells. This has been shown in earlier papers. This paper deals with the growth of strains of cells (Daucus carota and Haplopappus gracilis cells were used), washed and filtered free from the larger groups, on nutrient agar media in Petri dishes, thus simulating familiar microbiological technique. Each discrete member of a suspension is referred to as a “unit.” On the order of 1–10% of the separate units of a suspension may be induced to grow into viable colonies, depending on the strain and the conditions employed. Whereas at least 30% of the free single carrot cells were shown to be capable of division, only up to about 4% continued their growth to form macroscopically visible colonies when they were widely dispersed. Coconut milk promotes the growth of carrot cells into colonies. Both coconut milk and napthaleneacetic acid, which interact synergistically, arc required for the growth of Haplopappus cells. Various techniques which affect viability (the frequency with which units grow into colonies) were investigated. The viability of carrot units was found (1) to increase with their density on the plates; (2) to decrease upon washing the suspensions prior to plating; (3) to increase with increasing initial size; and (4) to decrease to a vanishingly low value in rigorously filtered suspensions which consist principally of single cells, although the single cells were found to grow with appreciable frequency when the larger units were also present; and (5) to increase dramatically (100-fold) when a rigorously filtered suspension was plated on a medium upon which pieces of growing cultured tissue were placed. Thus, the induction of growth in free cells is enhanced, even in an environment nutritionally optimal for the growth of the larger cell masses, by as yet unknown factors which are contributed by the cells themselves, or by adjacent cells or groups of cells. It is suggested that within a group of cells growing in culture, and perhaps also in the organized growing regions of intact plants, the dividing cells are nourished or stimulated by adjacent but less frequently dividing cells. The implications of these results are discussed.