Rate-limiting factors in leaf photosynthesis. I. Carbon fluxes in the calvin cycle

Rates of photosynthesis of spinach leaves were varied by varying light intensity and CO₂ concentration. Metabolism of the leaves was then arrested by freezing them in liquid nitrogen. Chloroplasts were isolated by a nonaqueous procedure. In the chloroplast fractions, levels of intermediates of the carbon reduction cycle were determined and considered in relation to the photosynthetic flux situation of the leaves at the time before freezing. During induction of photosynthesis, ribulose 1,5-bisphosphate levels increased in parallel with CO₂ fixation. In the steady state, a similar relation between ribulose 1,5-bisphosphate levels and CO₂ uptake was observed at light intensities between 0 and 50 W·m⁻². A further increase in light intensity increased CO₂ fixation rates but not ribulose 1,5-bisphosphate levels. Increasing the CO₂ concentration resulted in increased CO₂ uptake, whereas ribulose 1,5-bisphosphate levels decreased. Even under CO₂ saturation, ribulose 1,5-bisphosphate levels were about 100 nmol/mg chlorophyll corresponding to about 3.5 mM ribulose 1,5-bisphosphate in the chloroplast stroma. This suggests that even under CO₂ saturation, ribulose-1,5-bisphosphate carboxylase limits photosynhetic CO₂ uptake. Mass action ratios calculated from measured metabolite levels demonstrated that the thermodynamic gradient required for the regeneration of ribulose 1,5-bisphosphate from hexosephosphate and triosephosphate increased considerably as photosynthetic flux increased. Similar calculations revealed that the enzymatic apparatus responsible for the reduction of 3-phosphoglycerate to dihydroxyacetone phosphate is not displaced much from equilibrium even under maximum rates of photosynthesis at saturating CO₂. The same is true for aldolase. Fructose-1,6-bisphosphatase also did not limit Calvin cycle turnover. Only at very low light intensities and during the first minutes of the induction period was the ratio of fructose 1,6-bisphosphate to fructose 6-phosphate high. This observation was more readily explained in terms of fructose 1,6-bisphosphate binding to ribulose-1,5-bisphosphate carboxylase than by a rate limitation imposed by insufficient activation of fructose-1,6-bisphosphatase.     

Diurnal growth of tall fescue leaf blades : I. Spatial distribution of growth, deposition of water, and assimilate import in the elongation zone

Tall fescue leaf blades elongate at near constant rates during most of the light and dark periods of the diurnal cycle, with the dark rate being higher by 60 to 65%. Our objective was to determine relationships among diurnal rates of leaf elongation, deposition of water and deposition of dry matter (DM) into the elongation zone. Two separate experiments were conducted, both with a 15-hour photoperiod and constant 21°C at the growth zone. Increased rates of leaf elongation in darkness were due to proportionally increased rates of elongation of 4-millimeter segments of the elongation zone. Length of the total elongation zone was 30 millimeters in both light and darkness. The spatial distribution of water contents in the elongation zone varied little during the diurnal cycle. Thus, dark stimulation of leaf elongation rate (+65%) and of water deposition (+77%) into elongation zones were similar. Water content per unit leaf length increased by 50% between the basal and distal limits of the elongation zone, indicating that tissue also grew in the lateral and vertical dimensions. Longitudinal growth of tissue, however, allowed 5 to 7 times more water deposition into the elongation zone than growth in cross-sectional area. This relationship was similar in light and darkness. In both light and darkness net rates of DM deposition (microgram per millimeter leaf length per hour) increased from the zone of cell division towards the region of most active elongation, 10 to 15 millimeters from the ligule, then decreased towards the distal end of the elongation zone. Net DM deposition rates (microgram per hour) integrated over the 30-millimeter elongation zone were similar during light and darkness. Thus, DM in the elongation zone was diluted during darkness as a result of increased water deposition. Net DM deposition rates at and above the distal end of the elongation zone were clearly positive during the light, but were close to zero or negative in darkness. Thus, DM deposition into the elongation zone and the adjacent recently expanded tissue was differentially affected in the diurnal cycle, DM deposition occurred in both tissues in light, but was restricted to the elongation zone in darkness.     

NADP-malic enzyme from maize leaf: purification and properties.

NADP-malic enzyme (EC 1.1.1.40), which is involved in the photosynthetic C⁴ pathway, was isolated from maize leaf and purified to apparent homogeneity as judged by polyacrylamide gel electrophoresis. At the final step, chromatography on Blue-Sepharose, the enzyme had been purified approximately 80-fold from the initial crude extract and its specific activity was 101 μmol malate decarboxylated/mg protein/min at pH 8.4. The enzyme protein had a sedimentation coefficient (s_20,w) of 9.7 and molecular weight of 2.27 × 10⁵ in sucrose density gradient centrifugation, and molecular weight of 2.26 × 10⁵ calculated from sedimentation equilibrium analysis. The molecular weight of the monomeric form was determined to be 6.3 × 10⁴ by sodium dodecyl sulfate-polyacrylamide gel electrophoresis. In the pyruvate carboxylation reaction, HCO₃^? proved to be the active molecular species involved. With all other substrates at saturating concentration, the following kinetic constants were obtained: K_m (malate), 0.4 mm; K_m (NADP), 17.6 μm; K_m (Mg²⁺), 0.11 mm. The maize leaf malic enzyme was absolutely specific for NADP. The Arrhenius plot obtained from enzyme activity measurements was linear in a temperature range of 13 to 48 °C, and the activation energy was calculated to be 9500 cal/mol.     

NADP-malic enzyme from maize leaf: regulatory properties.

The regulatory properties of purified maize leaf NADP-malic enzyme (EC 1.1.1.40) were studied at three different pHs and the following results were obtained. (a) At pH 7.5 enzyme activity reaches a maximum at 0.4–0.8 mm malate depending on the Mg²⁺ concentration, and higher levels of malate result in marked substrate inhibition; with increasing pH the degree of substrate inhibition is reduced to where at pH 8.4 little or no inhibition is observed. (b) The inhibitory effect of malate is more pronounced at 1 mm Mg²⁺ than at 5–10 mm Mg²⁺ in the pH range of 7.5 to 8.4; a plot of enzyme activity vs Mg²⁺ concentration at 3 mm malate follows Michaelis-Menten kinetics at both pH 7.5 and 8.4; the apparent affinity of the enzyme for Mg²⁺ at pH 8.4 was threefold greater than that at pH 7.5. (c) The activity of NADP-malic enzyme decreases as the ratio of $\text{NADPH}\text{NADP}$ increases, and this effect is enhanced at lower pH. (d) Various α-keto acids including glyoxylate, oxaloacetate, and α-ketoglutarate inhibit NADP-malic enzyme activity, whereas HCO₃^?, pyruvate, and other organic acids, sugar phosphates, and amino acids have little or no effect on the activity of the enzyme. Based on these experimental findings, the regulatory properties of maize leaf NADP-malic enzyme are discussed with respect to its key role in net CO₂ fixation in maize bundle sheath chloroplasts during C₄ photosynthesis.     

伤根对玉米光合作用和水分利用效率的影响

1　引　　言在世界范围内 ,水资源的短缺日益受到人们的关注 ,农业水资源的高效利用已是世界农业亟待解决的主要问题 .因此 ,现代农业不应再单纯满足于高产 ,还应着眼于节约资源 ,提高水资源利用效率 .近年来 ,为了提高产量及水分利用效率 ,农业科技工作者在植物 水分方面做了大量的工作[1～ 3 ,5～ 7,9～ 12 ] .在人类的农业生产实践中 ,水稻插秧、幼苗移栽以及对作物的中耕等都可以使植株生长状况好转 ,作物产量提高 ,而这些措施都会对植物根系产生一定的影响 .对作物根系进行人为的伤害 ,也可能会改善作物生长状况 ,调节作物对有限土壤水…     

Diurnal fluctuations in cotton leaf carbon export, carbohydrate content, and sucrose synthesizing enzymes

In fully expanded leaves of greenhouse-grown cotton (Gossypium hirsutum L., cv Coker 100) plants, carbon export, starch accumulation rate, and carbon exchange rate exhibited different behavior during the light period. Starch accumulation rates were relatively constant during the light period, whereas carbon export rate was greater in the afternoon than in the morning even though the carbon exchange rate peaked about noon. Sucrose levels increased throughout the light period and dropped sharply with the onset of darkness; hexose levels were relatively constant except for a slight peak in the early morning. Sucrose synthase, usually thought to be a degradative enzyme, was found in unusually high activities in cotton leaf. Both sucrose synthase and sucrose phosphate synthetase activities were found to fluctuate diurnally in cotton leaves but with different rhythms. Diurnal fluctuations in the rate of sucrose export were generally aligned with sucrose phosphate synthase activity during the light period but not with sucrose synthase activity; neither enzyme activity correlated with carbon export during the dark. Cotton leaf sucrose phosphate synthase activity was sufficient to account for the observed carbon export rates; there is no need to invoke sucrose synthase as a synthetic enzyme in mature cotton leaves. During the dark a significant correlation was found between starch degradation rate and leaf carbon export. These results indicate that carbon partitioning in cotton leaf is somewhat independent of the carbon exchange rate and that leaf carbon export rate may be linked to sucrose formation and content during the light period and to starch breakdown in the dark.     

Diurnal changes in net uptake rate of nitrate are associated with changes in estimated export of carbohydrates to roots

The rate of NO3- uptake by soybean (Glycine max [L.] Merrill) roots generally declines during the night in association with progressive depletion of the nonstructural carbohydrate pool in the shoot as well as the concentration of carbohydrates in roots. To determine if NO3- uptake rate changes in response to variations in translocation rate of carbohydrates from shoot to roots per se or to carbohydrate status of the roots, the night period was interrupted with a low light level from incandescent lamps to alter the diurnal pattern of NO3- uptake by roots and export of carbohydrate from shoots of nonnodulated soybean. Depletion of NO3- from replenished, complete nutrient solutions containing 1 mM NO3- was measured by ion chromatography and rates of NO3- uptake were calculated. Changes in export of carbohydrates from shoot to roots during intervals of the night period were calculated as the differences between rates of disappearance in contents of nonstructural carbohydrates and their estimated rates of utilization in shoot respiration and growth. A positive, significant correlation occurred between changes in calculated rates of carbohydrate export from shoots and NO3- uptake rates. Conversely, there was no significant correlation between concentrations of nonstructural carbohydrates in roots and NO3- uptake rates. These results support the hypothesis that carbohydrate flux from shoot to roots has a direct role in regulation of nitrogen uptake by the whole plant.     

Diurnal changes in leaf gas exchange and chlorophyll fluorescence in tropical tree species with contrasting light requirements

Interspecific ecophysiological differences in response to different light environments are important to consider in regeneration behavior and forest dynamics. The diurnal changes in leaf gas exchange and chlorophyll fluorescence of two dipterocarps, Shorea leprosula (a high light-requiring) and Neobalanocarpus heimii (a low light-requiring), and a pioneer tree species (Macaranga gigantea) growing in open and gap sites were examined. In the open site, the maximum net photosynthetic rate (P_n), photosystem II (PSII) quantum yield (; F/Fm), and relative electron transport rate (r-ETR) through PSII at a given photosynthetic photon flux density (PPFD) was higher in S. leprosula and M. gigantea than in N. heimii, while non-photochemical quenching (NPQ) at a given PPFD was higher in N. heimii. The maximum values of net photosynthetic rate (P_n) in M. gigantea and S. leprosula was higher in the open site (8–11 mol m^–2 s^–1) than in the gap site (5 mol m^–2 s^–1), whereas that in N. heimii was lower in the open site (2 mol m^–2 s^–1) than in the gap site (4 mol m^–2 s^–1), indicating that N. heimii was less favorable to the open site. These data provide evidence to support the hypothesis that ecophysiological characteristics link with plants regeneration behavior and successional status. Although P_n and stomatal conductance decreased at midday in M. gigantea and S. leprosula in the open site, both r-ETR and leaf temperature remained unchanged. This indicates that stomatal closure rather than reduced photochemical capacity limited P_n in the daytime. Conversely, there was reduced r-ETR under high PPFD conditions in N. heimii in the open site, indicating reduced photochemical capacity. In the gap site, P_n increased in all leaves in the morning before exposure to direct sunlight, suggesting a relatively high use of diffuse light in the morning.     

断根对盐胁迫下玉米生长、光合及叶片抗氧化酶的影响

通过盆栽试验,以全根玉米为对照,研究了断根对不同浓度NaC1(无盐0、低盐0.2％、中盐0.4％、高盐0.6％)胁迫下2个玉米杂交种(郑单958、登海9)的生长、灌浆期光合及叶片抗氧化酶活性的影响.结果表明:低盐胁迫下,断根郑单958籽粒产量比全根玉米增加13.1％,登海9籽粒产量比全根玉米增加31.4％.拔节期,玉米生长受断根影响,其根及地上部干质量小于同盐度下的全根玉米,无盐和低盐条件下断根玉米生长恢复迅速.乳熟期,无盐和低盐条件下断根玉米的根及地上部干质量、叶面积、根总长、根总表面积、根系活力、叶绿素含量,以及穗位叶净光合速率、气孔导度、蒸腾速率、POD和CAT活性均显著高于全根玉米,而茎粗和穗位叶MDA含量小于全根玉米;中、高盐度胁迫对断根玉米影响较大,干质量、根系形态及光合指标均小于全根玉米,产量也略低于全根玉米.在整个生育期,断根对盐胁迫下玉米生长的影响与盐浓度有关,无盐和低盐浓度下促进玉米生长,中、高盐度则抑制生长.     

Diurnal changes in leaf water potential of rice,barley and wheat

The diurnal changes in leaf water potential of rice, barley and wheat corresponded to changes in air temperature. The soil moisture did not affect the diurnal changes in leaf water potential of rice and only slightly affected that of barley and wheat.     

超高产夏玉米田土壤微生物与土壤酶的动态变化

为揭示超高产夏玉米田（产量>15 000 kg·hm^-2）土壤微生物与土壤酶活性动态变化特性,在国家玉米工程技术研究中心（山东）试验场进行夏玉米生长季农田土壤微生物与土壤酶活性研究. 在连续3年产量15 000 kg·hm^-2以上的超高产夏玉米田中选择一块超高产田（HF, 产量为20 322 kg·hm^-2）与常规生产田（CF, 产量为8920.1 kg·hm^-2）进行对比分析,主要测定0～20 cm土层土壤细菌、真菌与放线菌数量及脲酶和转化酶活性. 结果表明：播种后超高产田与常规生产田土壤微生物（细菌、真菌与放线菌）数量均表现出先升高后下降的趋势,超高产田在玉米生长后期土壤微生物数量低于常规生产田,细菌与放线菌表现尤其明显,收获期超高产田B/F值（细菌与真菌数量比）比播种期高2.03倍,比常规生产田高3.02倍,常规生产田收获期与播种期的B/F值变化不显著;超高产田土壤脲酶活性在播种31 d（拔节期）后低于常规生产田,转化酶活性播种58 d（开花期）后快速下降,低于常规生产田.     

Flooding-induced changes in photosynthesis and oxidative status in maize plants

Maize plants (Zea mays L.) were subjected to soil flooding for 72, 96, and 120 h. A noticeable decrease in the rate of net photosynthesis (P_N) and the activity of ribulose-1,5-bisphosphate carboxylase (RuBPC, EC 4.1.1.39) were observed. The values of intercellular CO₂ concentrations (c_i) increased in all flooded plants without significant changes in stomatal conductance (g_s). The activity of phosphoenolpyruvate carboxylase (PEPC, EC 4.1.1.31) increased twofold 120 h after soil flooding. Flooding of maize plants led to a decrease in chlorophyll and protein levels and to slight increase of proline content. Flooded plants exhibited a large accumulation of leaf acidity. An increase in the values of some important parameters associated with oxidative stress, namely peroxides production, lipid peroxidation, and electrolyte leakage, confirmed the suggestion that root oxygen deficiency caused photooxidative damage in maize leaves.     

Influence of leaf age on photosynthesis, enzyme activity, and metabolite levels in wheat

The rate of photosynthesis under high light (1000 micromole quanta per square meter per second) and at 25°C was measured during development of the third leaf on wheat plants and compared with the activity of several photosynthetic enzymes and the level of metabolites. The rate of photosynthesis reached a maximum the 7th day after leaf emergence and declined thereafter. There was a high and significant correlation between the rate of photosynthesis per leaf area and the activities of the enzymes ribulose 5-phosphate kinase (r = 0.91), ribulose 1,5-bisphosphate (RuBP) carboxylase (r = 0.94), 3-phosphoglycerate (PGA) kinase (r = 0.82), and fructose 1,6-bisphosphatase (r = 0.80) per leaf area. There was not a significant correlation of photosynthesis rate with chlorophyll content. The rate of photosynthesis was strongly correlated with the level of PGA (r = 0.85) and inversely correlated with the level of triose phosphate (dihydroxyacetone phosphate and glyceraldehyde 3-phosphate) (r = 0.92). RuBP levels did not change much during leaf development; therefore photosynthesis rate was not correlated with the level of RuBP. The rate of photosynthesis was at a maximum when the ratio of PGA/triose phosphate was high, and when the ratio of RuBP/PGA was low. Although several enzymes change in parallel with leaf development, the metabolite changes suggest the greatest degree of control may be through RuBP carboxylase. The sucrose content of the leaf was highest under high rates of photosynthesis. There was no evidence that later in leaf development, photosynthesis (measured under high light and at 25°C) was limited by utilization of photosynthate.     

Growth, photosynthesis and assimilate partitioning in Lolium temulentum exposed to chilling temperatures

Vegetative plants of Lolium temulentum L, grown at 20°C with an 8 h photoperiod were transferred to either 5 or 2°C (8 h photoperiod) at 4th leaf maturity. Measurement of dry weight gain indicated a marked decline in relative growth rate below 5°C, with growth being reduced as much between 5 and 2°C as between 20 and 5°C. This reduction was not associated with increased mortality and was reversible if plants were returned to 20°C. Tissue explants from cold-treated plants retained the ability to extend if returned to 20°C Rates of extension in explants were less temperature sensitive than the relative growth rates observed in intact plants. Measurements of photosynthetic capacity, and of the patterns of accumulation of reserve carbohydrate in plants exposed to different temperatures, suggested that the inhibition of growth caused by chilling is not caused by an inability of the plants to fix sufficient carbon.     

Carbon dioxide exchange rates, ribulose bisphosphate carboxylase/oxygenase and phosphoenolpyruvate carboxylase activities, and kernel growth characteristics of maize

Four high-yield-potential maize hybrids (FS854, CB596 x LH38, B73 x LH38, and B73 x Mo17) and four inbred lines (LH38, CB59G, Mo17, and B73) were grown in the field to study traits associated with leaf area duration (LAD) and the relationship between LAD and kernel growth characters. Based on decline in chlorophyll, leaf N concentration, CO(2) exchange rate, and ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPCase) activities, the hybrid B73 x Mo17 had a significantly shorter LAD than the other three hybrids. The shorter LAD was not due to maturity because B73 x Mo17 is in a maturity class similar to the other hybrids except CB59G x LH38, which is approximately 1 week earlier. At the time of grain maturity, leaves of B73 x Mo17 had lost all chlorophyll and CO(2) exchange and carboxylase activities. The other three hybrids, however, retained green leaves which still had 20% of the maximum CO(2) exchange rate. In addition, B73 x Mo17 remobilized leaf N more extensively. For all hybrids, declines in CO(2) exchange were closely correlated with declines in PEPCase activity, whereas the relationship between CO(2) exchange and Rubisco activity was weak. Responses of the inbred lines predicted, to some extent, physiological characteristics of the hybrids. CB59G and LH38 both had a longer LAD than either B73 or Mo17 as judged by decline in chlorophyll, leaf N, CO(2) exchange rate, and Rubisco and PEPCase activities. With the exception of B73 x LH38, kernel growth characteristics of the hybrids were related to LAD. Effective filling period (EFP) measured in days was 32.9, 31.5, 30.8, and 30.4 for FS854, CB59G x LH38, B73 x LH38, and B73 x Mo17, respectively. For FS854 and CB59G x LH38, the longer EFP was associated with a larger kernel weight. These data suggested that late season photoassimilate resulting from longer LAD could be utilized by the kernels of these two hybrids. For B73 x Mo17, the shorter LAD and EFP was associated with a kernel dry matter accumulation rate (10.1 milligrams per kernel per day) which was significantly higher than for the other three hybrids. Thus, the more rapid leaf senescence of B73 x Mo17 appeared to be coordinated with efficient leaf N remobilization and a relatively short grain-filling period characterized by rapid kernel dry matter accumulation.     

低温弱光下水杨酸对黄瓜幼苗光合作用及抗氧化酶活性的影响

为了探讨低温弱光下水杨酸（SA）对黄瓜光合功能的调控作用,以‘津优3号’黄瓜幼苗为试材,叶面喷施不同浓度的SA溶液,研究低温弱光下黄瓜幼苗气体交换参数、光化学效率、MDA含量及抗氧化酶活性的变化.结果表明：低温弱光胁迫使黄瓜幼苗叶片的光合速率（P_n）、气孔导度（G_s）、蒸腾速率（T_r）、PSⅡ光下实际光化学效率（Φ_PSⅡ）及暗下最大光化学效率（F_v/F_m）明显降低,胞间CO₂浓度（C_i）显著升高,说明低温弱光下黄瓜幼苗P_n下降的主要原因是非气孔限制;低温弱光还可引起黄瓜幼苗丙二醛（MDA）含量增加,超氧化物歧化酶（SOD）活性升高,过氧化氢酶（CAT）活性降低,过氧化物酶（POD）活性先升高后降低.而胁迫前用0.5～2.5 mmol·L^-1 SA预处理幼苗,其叶片的P_n、G_s、T_r、Φ_PSⅡ、F_v/F_m及SOD、POD和CAT活性与CK（水预处理）相比均有不同程度的提高,C_i和MDA含量有所降低.表明SA可有效调控低温弱光下黄瓜幼苗叶片的光合功能,提高其低温弱光耐性,其适宜浓度为1 mmol·L^-1.     

Biochemical basis for effects of k-deficiency on assimilate export rate and accumulation of soluble sugars in soybean leaves

The effects of K-deficiency on carbon exchange rates (CER), photosynthate partitioning, export rate, and activities of key enzymes involved in sucrose metabolism were studied in soybean (Glycine max [L.] Merr.) leaves. The different parameters were monitored in mature leaves that had expanded prior to, or during, imposition of a complete K-deficiency (plants received K-free nutrition solution). In general, recently expanded leaves had the highest concentration of K, and imposition of K-stress at any stage of leaf expansion resulted in decreased K concentrations relative to control plants (10 millimolar K). A reduction in CER, relative to control plants, was only observed in leaves that expanded during the K-stress. Stomatal conductance also declined, but this was not the primary cause of the decrease in carbon fixation because internal CO₂ concentration was unaffected by K-stress. Assimilate export rate from K-deficient leaves was reduced but relative export, calculated as a percentage of CER, was similar to control leaves. Over all the data, export rate was correlated positively with both CER and activity of sucrose phosphate synthase in leaf extracts. K-deficient leaves had higher concentrations of sucrose and hexose sugars. Accumulation of hexose sugars was associated with increased activities of acid invertase. Neutral invertase activity was low and unaffected by K-nutrition. It is concluded that decreased rates of assimilate export are associated with decreased activities of sucrose phosphate synthase, a key enzyme involved in sucrose formation, and that accumulation of hexose sugars may occur because of increased hydrolysis of sucrose in K-deficient leaves.     

Developmental changes and regional distribution of phospholipase D and base exchange enzyme activities in rat brain

Phospholipase D (PL-D) activity per mg protein of whole homogenate increased 5.1 fold between Embryonic (E) day 17 and Postpartum (P) day 14 and slightly decreased by P 30 days. This was due to the decrease of PL-D activity of the P₂ fraction. The PL-D activity of P₂ and P₃ fractions increased 11.2 and 6.1 fold respectively between E 17 and P 14. The 3 base exchange enzyme (BEE) activities per mg protein of whole homogenate increased up to P 14 or P 21 and then decreased. This decrease was greater in the P₂ fraction and the P₃ fraction increased after P14. Brains from 1 day to 25 month old rats were dissected into 7 separate regions and both PL-D and BEE activities were measured. In adult rats, the hippocampus and hypothalamus had the highest PL-D activities while medulla+pons and cerebellum had the lowest PL-D activities. The developmental patterns of 5 regions except for hippocampus and hypothalamus were similar. PL-D activity in the hippocampus was maximum at P 7 followed by a steep decrease till P30 suggesting that the PL-D activity of the hypothalamus develops later and that of the hippocampus develops earlier than any other region. The distributions of BEE activities were quite different from those of PL-D activities. In adult rats, the cerebellum had the highest activity while the striatum and medulla+pons had the lowest. The BEE activities of cerebellum were lowest at P 1 and showed steep increase during the next 2 weeks.To whom to address reprint request are to be sent.