A Quantitative Analysis of the Uptake of Carbon and of the Supply of 14C-labelled Assimilates to Areas of Meristematic Growth in Lolium temulentum

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby leaves on the main shoot to terminal meristem, stem, tillers,and roots was conducted during parallel periods of reproductiveand vegetative development in Lolium temulentum. The initial rate of entry of carbon into the shoot varied withthe area and photosynthetic efficiency of the assimilating leaf.Subsequently, respiratory losses of carbon during translocationand incorporation of assimilate at the site of utilization alsovaried. The combined effect of these differences resulted inthe supply of recently assimilated carbon being twofold greaterin reproductive shoots than in vegetative shoots, while withinshoots the carbon supply of the youngest fully expanded leafranged from four-or five-fold greater than the oldest leaf inyoung shoots, to two-or three-fold greater in older shoots.In both reproductive and vegetative shoots, the two or threeyoungest leaves thus dominated the supply of carbon for meristematicgrowth. Meristematic tissue in expanding leaves and leaf primordia atthe terminal meristem of the vegetative shoot received 18–27per cent of the total shoot carbon. This meristem utilized aboutthe same proportion of shoot carbon when it developed into aninflorescence, indicating no major change in the level of meristematicactivity. The proportion of shoot carbon utilized in stem growthincreased as both reproductive and vegetative shoots aged; thisincreased meristematic activity in stem internodes was accompaniedby reduced export of carbon to roots, which received less than10 per cent of the shoot carbon when the experiments ended.The main shoot translocated 20–30 per cent of its recentlyassimilated carbon to developing and rooted tillers, which assinks for carbon were thus as important as the terminal meristemand stem. This outward flow of carbon continued relatively uncheckedwhen donor and receptor shoots developed inflorescences.     

Effects of Temperature on the Distribution of 14C-labelled Assimilates in the Flowering Shoot of Carnation

The pattern of distribution of ¹⁴C-labelled assimilates translocatedfrom a leaf on the flower stem of carnation was found to varywith both the ambient air temperature and the localized temperatureof the flower bud. A high bud temperature increased the proportionofassimilates moving into the floral tissues while a low budtemperature increased the proportion accumulating in the stemabove the source leaf. When the air temperature was raised independentlyof the bud temperature, the stem gained assimilates at the expenseof the flower, but if both air temperature and bud temperaturewere raised together, effects of bud temperature predominatedand movement of assimilates into the flower was promoted. Therole of the flower in mediating effects of temperature is discussedwith reference to the distribution of invertase activity inthe shoot.     

Hormonal Regulation of the Distribution of 14C-labelled Assimilates in the Flowering Shoot of Carnation

Evidence is presented that growth substances have a role indirecting the movement of assimilates towards the developingflowers of carnation. Removal of the terminal flower resultedin decreased upward movement of ¹⁴C-labelled assimilates froma source leaf on the flowering shoot. Applications of indole-3yl-aceticacid or gibberellic acid to the stump of the flower stem partiallyrestored the pattern of movement to that found in the intactshoot. Two gibberellin-like substances and one auxin-like substancewere detected in extracts of the floral tissues. Levels of growthsubstances appeared to be higher in the flower than elsewherein the flowering shoot.     

Independent translocation of 14C-labelled assimilates and of the floral stimulus in Lolium temulentum

Summary It is widely accepted that the floral stimulus produced in leaves is carried to the shoot apex passively in the phloem with the assimilate stream. Three kinds of evidence presented here suggest that the floral stimulus moves independently of the assimilates.Simultaneous determination of the velocities of translocation out of the seventh leaf blade, in comparable plants under the same conditions, yielded estimates of 1–2.4 cm/hr for the floral stimulus, and 77–105 cm/hr for ¹⁴C-labelled assimilates.The effect of the size of the seventh leaf on its ability to export assimilates or to initiate flowering was quite different. Leaves with only 14–26% of their final blade area emerged exported little assimilate, yet were highly active in inducing flowering.The effect of DCMU applications at a range of concentrations on the translocation of assimilates was quite different from their effect on the flowering response.     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.): 3. Partition of Assimilates between Root and Shoot during the Transition from Vegetative to Reproductive Growth

The partition of ¹⁴C labelled current assimilates to root insimulated swards of Lolium perenne cv. S24 was measured duringthe transition from vegetative growth in autumn to reproductivegrowth in spring under close to natural conditions of lightand temperature. Assimilate partitioning was also measured in‘established’ swards cut three times during thegrowing season and in vegetative ‘seedling’ swardsgrowing in autumn and in spring. All measurements were madewhen the swards had achieved more than 90 per cent light interception,and all swards were abundantly supplied with water and mineralnutrients. During autumn there was a gradual decrease in the proportionof assimilates partitioned to the roots in both the ‘established’and the ‘seedling’ swards. In the established swards,partition to roots was low over winter, increased during earlyspring, but decreased dramatically, later in the spring, whenstem elongation began. In contrast, in the unvernalized vegatativeseedling swards in spring, partition to roots remained high. The seasonal pattern of assimilate partitioning is consideredin relation to changes in the natural environment and the rateat which the crop fixed carbon in photosynthesis. A decreasein the proportion of assimilates partitioned to roots duringlate spring was significant in increasing the production ofshoot at that time but seasonal differences in partition contributedvery little to the marked differences in shoot growth betweenthe spring and autumn crop. Lolium perenne L., perennial ryegrass, partition of assimilates, flowering     

The Effects of Leaf Age and Senescence on the Distribution of Carbon in Lolium temulentum

Assimilate distribution in leaves of Lolium temulentum was establishedby root absorption of [¹⁴C]sucrose and after exposure to ¹⁴CO₂.Age determined the amount of carbon assimilated, with more labelbeing incorporated during expansion than at maturity. Duringsenescence ¹⁴C assimilation was much lower. Ethanol-solubleextracts from various tissues of root-labelled plants containedmost of the radioactivity chiefly in basic and acidic compounds.The neutral fraction was composed predominantly of sucrose. Sucrose was comparably labelled in leaves from plants fed equalamounts of either [¹⁴C]sucrose, glucose, or fructose and onlytraces of labelled monosaccharides appeared in extracts. Radioactive sucrose was translocated rapidly from mature leaveswhereas, in the expanding leaf, carbon incorporation was directedtowards growth and the greater proportion of label present atligule formation was in ethanol-insoluble material. Induced senescence, of a mature leaf fed during expansion, produceda rapid loss from the pool of insoluble ¹⁴C. This was accompaniedby a reduction in the contents of chlorophyll and soluble proteinand an accumulation of amino acids. The onset of senescencecaused changes in leaf sugar levels which were correlated withincreased rates of respiration.     

The Influence of Plant Water Deficit on Distribution of 14C-labelled Assimilates in Cacao Seedlings

The relationship between plant water status and distributionof ¹⁴C-labelled assimilates in cacao (Theobroma cacao L.) wasevaluated after ¹⁴CO₂ pulse labelling leaves of seedlings subjectedto varying levels of water deficiency. The proportion of ¹⁴Cexported by source leaves was strongly affected by seedlingwater status. An increasing proportion of labelled assimilatesremained in source leaves at both 24-h and 72-h harvests aswater stress intensity increased. Water stress reduced the distributionof exported label to leaves and to the expanding flush in particularbut increased the proportion of label in stems and roots. Theresults suggest that current photoassimilates may be temporarilystored in source leaves and stems of cacao seedlings duringperiods of plant water deficit. The stress-induced changes inpartitioning of labelled carbon were in concordance with changesin shoot to root biomass ratios, which was likely due to greaterreduction in growth of above-ground organs to that of roots. Theobroma cacao L, assimilate partitioning, cacao, ¹⁴C-photoassimilate, water stress, water potential     

Kinetics of Leaf Growth in Lolium temulentum at Optimal and Chilling Temperatures

Lolium temulentum plants were grown at 20 °C, under an 8-hdaylength, in a controlled-environment chamber, and the kineticsof leaf expansion were observed by measuring the movement ofan optical grid attached to the fourth leaf. The leaf emerged23–24 d after sowing and was fully expanded 9–10d later. Extension rate was maximal between the second and fifthdays after emergence and declined markedly thereafter. Duringthe rapid growth phase the rate of elongation exhibited a distinctdiurnal rhythm, fluctuating between 1.9 to 2.3 mm h^–1in the light period, and 1.3 to 1.7 mm h^–1 in the dark.A circadian oscillation with a period of about 27 h was observedin leaves elongating in continuous darkness. When plants weretransferred to 5 °C soon after emergence of the fourth leafthere was an immediate reduction in rate of growth to about22 per cent of the rate at 20 °C: the Q₁₀ for the mean elongationrate in the range 20–5 °C was 3.7. When plants weretransferred from 20 to 2 °C at fourth leaf emergence, meanextension rate declined to less than 5 per cent, correspondingto a Q₁₀ in the range 5–2 °C of more than 300. Furthermore,growth at 2 °C was confined almost entirely to the darkphase of the photoperiod cycle. The responsive tissue was shownto be a small area of expanding leafless than 1.5 cm above theshoot apex and the possible mechanisms underlying low temperatureeffects in this region are discussed. Lolium temulentum L., leaf growth, auxanometer, low temperature, diurnal rhythm     

菊花不同生长阶段不同器官POD和EST同工酶比较

采用过氧化物酶(POD)、酯酶(EST)2个酶系统的12个同工酶位点,分析了4个菊花品种营养生长和生殖生长阶段不同器官(嫩叶、老叶、嫩茎、木质化茎)的同工酶变化.结果表明:(1)4个品种共有16种POD酶带,15种EST酶带;(2)菊花的POD和EST具有组织特异性和阶段特异性,其中以嫩叶的酶带最多,其次为老叶,再次为嫩茎,而木质化茎的酶带最少;(3)与生殖生长阶段相比,营养生长阶段的POD酶带更清晰,更整齐,分离更好,但生殖生长阶段的EST同工酶比营养生长阶段的更清晰;(4)营养生长阶段的嫩叶最适合用于菊花POD同工酶分析,而EST同工酶研究宜取生殖生长时期的嫩叶.     

A Catastrophe Model for the Switch from Vegetative to Reproductive Growth in the Shoot Apex

A mathematical model is constructed to describe the morphopneticswitch that occurs when a vegetative plant apex becomes reproductive.The cusp equation from catastrophe theory is modified, and isused to relate primordial size at initiation to apex size. Theresulting equation may be viewed as an equation of state definingthe allowed organizational modes of the shoot apex. The modelsimulates the growth of the apex from the vetative stage toearly reproductive growth, and makes reasonable predictionsabout apex size and growth rate, primordial sizes, and the lengthsof the plastochron. flowering, mathematical model, catastrophe theory, shoot apex     

Effects of Leaf Infections by Septoria nodorum Berk. on the Translocation of 14C-labelled Assimilates in Spring Wheat

The influence of infection with Septoria nodorum of leaves belowthe flag leaf on the translocation of ¹⁴C-labelled assimilatesin wheat was followed. In the vegetative phase export of assimilatesfrom a single infected leaf was reduced, but export from a healthyleaf on a heavily infected plant was increased. During the reproductivephase export from leaves was not affected by disease. Heavyleaf infection had little effect on the patterns of distributionof export especially during reproductive growth when only changesin the proportion of assimilates in leaf sheaths and tillerstumps were found. Distribution of export from a healthy flagleaf on an otherwise heavily infected plant was unaltered. Duringvegetative growth changes in the distribution of assimilateswere more marked, the greatest changes occurring when a singleinfected leaf on a healthy plant was exposed to ¹⁴CO₂.     

The Sugars Concerned in the Tangential Movement of 14C-labelled Assimilates in Willow

Analysis of bark sectors through which tangential movement hadoccurred showed that sucrose was present in greater quantitiesthan glucose or fructose. No evidence could be gained from suchanalyses, however, as to which of these three sugars was themobile species. The use of ¹⁴CO₂ supplied to the leaves, revealed that after24 hours of tangential movement, the greatest proportion ofthe activity in the bark sectors was present in sucrose. Byprogressively reducing the time during which tangential movementwas allowed to occur in these tracer experiments, a patternof results was obtained which completely supports the assumptionthat ¹⁴C-labelled assimilates are transported in a tangentialmanner round the stem as sucrose. The presence of activity inglucose and fructose is merely due to their production fromlabelled sucrose.     

Changes in the Pattern of Cell Division in the Shoot and Root Meristems of Lolium perenne during the Transition from Vegetative to Floral Growth

For Lolium perenne cv. Cropper, a system which resulted in 100%flowering comprised 90 short days (SD) at 4 ?C (vernalization)and 30 SD at 18 ?C followed by 8 long days (LD). The mitoticindex and G1 and G2 percentages were measured in the shoot androot apices of plants following 2, 5 or 8 LD and in SD controlssampled at the beginning and end of induction. Identical measurementswere made in plants given 48 SD at 18 ?C followed by 2, 5 or8 LD; plants remained vegetative in response to this treatmentlacking vernalization. Significant increases in both mitoticindex and meristem size occurred in the shoot apex in LD followingthe vernalizing, but not the non-vernalizing, treatment. A clusterof mitoses in the apical dome of the shoot apex was unique tothe vernalized plants given 5 or 8 LD. However, an increasein root meristem size occurred regardless of vernalization,but a significant increase in the mitotic index was limitedto vernalized plants given 5 or 8 LD. Whilst the vernalization-LDtreatment resulted in an increase in the G2 percentage in theshoot apex following 2, 5 or 8 LD, no such alteration was observedin the root meristem. Thus, the changes to the cell cycle whichcorrelated with flowering were increased mitotic indices andG2 percentages in the shoot apex at each sampling time and increasedmitotic indices in the root apex following 5 and 8 LD. Key words: Cell division, flowering, Lolium perenne L.     

The Influence of Defoliation on the Distribution of Assimilates in Lolium multiflorum Lam

Autoradiographs were made of plants of Lolium multiflorum Lam.after ¹⁴CO₂ had been fixed by selected leaves. The results showedthat labelled compounds were not translocated to other tillersbut were moved to the whole root system. This pattern of distributionwas changed when all or some of the tillers on the plant weredefoliated. Where a single undefoliated tiller remained, itinitially supplied the cut tillers with ¹⁴C-containing products,thus reintegrating a system of apparently independent tillers.When all the tillers were partially defoliated, labelled compoundswere no longer translocated to the root system. A further experimentsuggested that root reserves were not mobilized for regrowthfollowing defoliation. These results are discussed in termsof the integration of a grass plant in the vegetative state.     

Transport of 14C-labelled Assimilates and 32P-labelled Phosphate in Salix viminalis in Relation to Phyllotaxis and Leaf Age

Autoradiographic techniques have been used to investigate thetransport of ¹⁴C-assimilates and ³²P-phosphate in relation tophyllotaxis and age of leaves in Salix viminalis. The leavesborne on the young shoots of S. viminalis are arranged in fiveorthostichies. Three of these orthostichies share one main transportchannel, a second channel being shared by the leaves on theremaining two orthostichies. Where two orthostichies are separatedby an angular distance of 72°, transport may occur betweenleaves on these orthostichies. In addition to the phyllotaxis of the leafy shoot, it has beendemonstrated that the age of the leaf is a factor which determinesthe distribution of materials between leaves.     

Patterns of 14C-labelled Assimilate Partitioning in Red and White Clover During Vegetative Growth

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby the expanded leaves on the primary shoot to growing leaves,stem, lateral shoots (branches or stolons) and roots in redand white clover was conducted during vegetative growth. Stem growth of the primary shoot was inhibited in both cloversand utilized no energy resources. The growing leaves at theprimary shoot apex of white clover imported 4 per cent of theshoot's assimilate compared with 10 per cent in red clover.At the basal end of the primary shoot, the tap root of whiteclover imported 16 per cent of the shoot's assimilate comparedwith 22 per cent in red clover. Branches in red clover and stolonsin white clover were by far the largest sinks for primary shootassimilate, importing 39 per cent and 63 per cent of the labelledassimilate, respectively. Analyses of the translocation of assimilate from individualprimary shoot leaves demonstrated that in both clovers olderleaves exported more of their assimilate to branches or stolons,whereas younger leaves exported more of their assimilate toroots, and possibly in white clover, to growing leaves at thetip of the shoot. Of the labelled assimilate exported to branchesor stolons, each primary shoot leaf exported preferentiallyto the branch or stolon in its own axil, but in addition exportedsubstantial quantities of assimilate to all other axillary shoots,particularly those arising from basal axils where the subtendingleaf had died. Trifolium repens, Trifolium pratense, red clover, white clover, assimilate partitioning, perennation     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.). 1. Response of Leaf Extension to Temperature during the Transition from Vegetative to Reproductive Growth

The potential for leaf extension of plants of Lolium perennecv S24 growing in small artificial communities under naturalconditions was measured as the plants progressed from the vegetativeto the reproductive state In two consecutive years, 1975 and 1976, ‘simulated swards’were sown in autumn and overwintered in an open, unheated glasshouseIndividual swards from the batch sown in 1975 were brought into a growth room on two occasions in spring 1976 to measuretheir potential for leaf extension at a range of temperatures(5–20 °C) Individual swards from the batch sown inautumn 1976 were brought in to the growth room on 15 occasionsbetween November 1976 and May 1977 and their potential for leafextension was measured at a single temperature of 15 °CFrequent dissections were made in both years to describe changesin the developing apex. The potential for leaf extension at 15 °C decreased fromc 17 mm day^–1 in November to c 10 mm day^–1 in mid-winter.In January, the potential rapidly increased threefold to reach30mm day^–1 by mid February The increase began coincidentwith the earliest stages of floral initiation and was completedby the time of double-ridge formation Spring-grown vegetativeplants, however, showed potential rates of < 20 mm day^–1at 15 °C The results are discussed in relation to reproductive developmentand to changes in the carbohydrate strategy of the plants inearly spring Lolium perenne L perennial ryegrass, leaf extension, temperature response     

小麦果皮绿色层中^1^4C标记同化物在胚珠内的分配与定位

小麦离体颖果饲喂~14CO_2后,经冷冻取代,于膜显微放射性自显影,追踪与检查了包括可溶性糖在内的~14C标记同化物在胚珠内的分配与定位。颖果绿色层细胞的高光合效率不仅保证了同化物的即时输出,而且还可以淀粉形式暂贮在叶绿体内。绿色层光合产物主要向腹沟处汇集,通过该处的维管束与珠心突起向胚乳方向迁移。~14C标记同化物以逆浓差梯度方式从珠心向珠心—胚乳交界处质外体空间输出并在胚乳内呈不均匀分布和富集于原胚端活跃增殖生长的区段内。绿色层同化物以多种方式进入原胚,主要以可溶态形式存在。同化物可能通过珠孔直接就近为原胚吸收。     

Preliminary Experiments on the Effect of Temperature on the Movement of 14C-labelled Assimilates through the Phloem of Willow

Experiments were performed on both young shoots of willow (3–5weeks old) and mature stems which had been growing for a periodof 2–3 years. ¹⁴CO₂ was supplied to the leaves, and themass transport of the labelled assimilates through a portionof the stem enclosed in a temperature-controlled jacket wasmeasured by determining the slope of the rise in the activityof huneydew collected from a colony of the aphid Tuberolachnussalignus (Gmelin). It was found that there was a marked difference in the behaviourof the young shoots and mature stems. In the former, a fallin temperature slowed the transport, whilst in the latter acomparable fall in temperature increased the transport. Possiblecauses for these effects have been suggested, and the resultsare discussed in relation to published work on temperature effects.