Metabolism in living systems is an integrated network of interacting processes directed toward an increase or conservation of the nonequilibrium. This is achieved by generating gradients of concentration, biosynthesis, or resynthesis of various compounds. In addition to integrity, metabolic processes are characterized by specialization, thereby giving rise to the multicomponent structure of metabolism. It was shown in the preceding work [1] that the structure of metabolic reactions in plant leaves consists of four components. It was also shown that one to four components of integral metabolism ( R s ) can be observed in plant leaves in accordance with their physiological state. The scheme of the four-component structure of plant metabolism is shown in Fig. 1. In this scheme, R b is the metabolic component maintaining the basic (functionally inactive) state of a native leaf. Because the leaf energy in this state is spent only to maintain its intactness, the structure of the leaf metabolism contains only the R b component. The ability to fulfil other functions is achieved during leaf activation. The activation is a structural rearrangement causing an increase in the membrane permeability, a decrease in the thermal stability of leaf structures, and other nonequilibrium processes. Therefore, an additional increase in the efficiency of sustaining metabolism and the corresponding energy expenditure are required to maintain activated leaves in the native state [1]. This additional increase in the efficiency of sustaining metabolism corresponds to the component R a . Therefore, R a is the metabolic component maintaining the leaf in the functionally active state. Therefore, supportive metabolism ( R m ) may contain one ( R b ) or two ( R b + R a ) components. R f and R g are the functional and growth components of leaf metabolism, respectively. R g is a component of R s only during plant growth. This component of leaf metabolism corresponds to the de novo biosynthesis. The other growthsupporting processes (primarily, transport processes) are regarded as auxiliary. Therefore, they were included in R f . Thus, R g and R f are coupled with one another, and R f is also coupled with R a . Indeed, as noted above, R a is the metabolic component maintaining the leaf in the functionally active state (Fig. 1). The temperature dependence of leaf metabolism has not yet been analyzed in terms of the four-component structure of plant leaf metabolism. The literature describes mainly the components R s , R f , and R g . The component R a has been isolated only recently, and there are few literature data on this component thus far. The temperature coefficient ( Q ) of the basic metabolism ( R b ) was calculated in the preceding work [2] only within the temperature range from 10 to 30 ie . Therefore, experimental study of temperature dependence of components R b and R a and comparative analysis of temperature dependences of the four components of leaf metabolism were the goals of this work. The literature data on the temperature dependence of photosynthesis [3], transport, and growth [4] were used to analyze the temperature dependence of the R f and R g components of leaf metabolism. Leaves of two plant species with contrasting thermal resistance were used: the thermophilous maize ( Zea mays L.) and the cold-resistant cabbage ( Brassica oleracia L.). Plants were grown in the field. Apparently healthy, separated, and mature leaves (i.e., leaves that did not grow any longer) were studied. After thick leaf ribs had been removed, leaf plates were cut into segments. The segment areas in cabbage and maize leaves were about 4 and 12 cm 2 , respectively. The leaf segments were mixed to obtain the total population, from which elementary samples (repetitions) were randomly taken. The total populations for the cabbage and maize leaves were 80 and 30 segments, respectively. The samples were incubated for 20 h at 25ie in a dark thermostat at an air humidity of 100%. Because metabolic reactions of mature leaves in situ do not contain the component R g , whereas the components R a and R f were eliminated during incubation [2, 5], metabolism of incubated leaves was characterized by a single component, R b . When incubation was over, 50% of the samples were activated by 20-min exposure to an ambient illuminance of 30 klx in a humid atmosphere containing no CO 2 . The lack of CO 2 inhibited biosynthesis of GENERAL BIOLOGY
Read full abstract