Dynamics of non-structural carbohydrates in developing leaves, bracts and floral buds of cotton

Development of cotton (Gossypium hirsutum L.) squares (i.e. floral buds with bracts) is fundamental for yield formation. A 2-year field study was conducted to determine dry weight (DW) accumulations of cotton leaves, floral bracts and floral buds, and the changes in concentrations of non-structural carbohydrates (hexoses, sucrose and starch) in these tissues during square ontogeny as affected by fruiting positions within the plant canopy. During square development, DW accumulation of a subtending sympodial leaf and floral bracts followed a sigmoid growth curve with increasing square age, whereas the DW increase of a floral bud followed an exponential curve. Main-stem node (Node 8, 10 or 12) and branch position (proximal vs. distal) within a plant canopy significantly affected DW accumulations of the leaf, bracts and floral bud. Starch was the dominant non-structural carbohydrate in the three tissues, accounting for more than 65% of total non-structural carbohydrates (TNC). Subtending leaf TNC increased as square age increased. The bracts exhibited a smaller change in TNC than leaves. Non-structural carbohydrate concentration was the lowest in 10-day-old floral buds, and had little change during the first 15 days of square development. Within 5 days prior to anthesis, the floral-bud TNC increased dramatically, tripling at the time of floral anthesis compared with 15-day-old floral buds. Square age and fruiting position significantly affected non-structural carbohydrate concentrations of subtending leaves, bracts, and floral buds. The correlation did not exist between final boll retention and non-structural carbohydrate concentrations of floral buds at different fruiting positions under normal growth conditions. The pattern of floral-bud non-structural carbohydrates during square ontogeny suggests that major events in carbohydrate metabolism occur just prior to anthesis.     

Water use efficiency as a function of leaf age and position within the cotton canopy

The gas exchange properties of whole plant canopies are an integral part of crop productivity and have attracted much attention in recent years. However, insufficient information exists on the coordination of transpiration and CO₂ uptake for individual leaves during the growing season. Single-leaf determinations of net photosynthesis (Pn), transpiration (E) and water use efficiency (WUE) for field-grown cotton (Gossypium hirsutum L.) leaves were recorded during a 2-year field study. Measurements were made at 3 to 4 day intervals on the main-stem and first three sympodial leaves at main-stem node 10 from their unfolding through senescence. Results indicated that all gas exchange parameters changed with individual main-stem and sympodial leaf age. Values of Pn, E and WUE followed a rise and fall pattern with maximum rates achieved at a leaf age of 18 to 20 days. While no significant position effects were observed for Pn, main-stem and sympodial leaves did differ in E and WUE particularly as leaves aged beyond 40 days. For a given leaf age, the main-stem leaf had a significantly lower WUE than the three sympodial leaves. WUE's for the main-stem and three sympodial leaves between the ages of 41 to 50 days were 0.85, 1.30, 1.36 and 1.95 μmol CO₂ mmol⁻¹ H₂O, respectively. The mechanisms which mediated leaf positional differences for WUE were not strictly related to changes in stomatal conductance (g_s·H₂O) since decreases in g_s·H₂O with leaf age were similar for the four leaves. However, significantly different radiant environments with distance along the fruiting branch did indicate the possible involvement of mutual leaf shading in determining WUE. The significance of these findings are presented in relation to light competition within the plant canopy during development.     

Influence of Potassium Deficiency on Photosynthesis,Chlorophyll Content,and Chloroplast Ultrastructure of Cotton Plants

In cotton (Gossypium hirsutum L.) grown in controlled-environment growth chamber the effects of K deficiency during floral bud development on leaf photosynthesis, contents of chlorophyll (Chl) and nonstructural saccharides, leaf anatomy, chloroplast ultrastructure, and plant dry matter accumulation were studied. After cotton plants received 35-d K-free nutrient solution at the early square stage, net photosynthetic rate (P _N) of the uppermost fully expanded main-stem leaves was only 23 % of the control plants receiving a full K supply. Decreased leaf P _N of K-deficient cotton was mainly associated with dramatically low Chl content, poor chloroplast ultrastructure, and restricted saccharide translocation, rather than limited stomata conductance in K-deficient leaves. Accumulation of sucrose in leaves of K-deficient plants might be associated with reduced entry of sucrose into the transport pool or decreased phloem loading. K deficiency during squaring also dramatically reduced leaf area and dry matter accumulation, and affected assimilate partitioning among plant tissues.     

Photosynthesis of bract and its contribution to seed maturity inCarpinus laxiflora

Light saturated net photosynthesis was measured in bracts and leaves ofCarpinus laxiflora, the major species in secondary forests in cool and intermediate temperate zones in Japan. The maximum net photosynthesis of leaves and bracts was essentially constant from May to early August and decreased gradually thereafter. For bracts, it was 3.2 μmol m⁻²s⁻¹, approximately half that for the leaves. The photosynthesis of bracts would thus appear to contribute significantly to seed maturity. The estimated production of bract based on the photosynthesis would make seeds (3 mg dry weight) mature for 37 days, assuming all photosynthate of the bracts to have been distributed in the seeds only. This was quite consistent with the growth curve for the seeds. A mast year phenomenon is discussed in relation to bract photosynthesis and leaf number.     

Photosynthetic and Respiratory Activity of Fruiting Forms within the Cotton Canopy

The supply of photosynthates by leaves for reproductive development in cotton (Gossypium hirsutum L.) has been extensively studied. However, the contribution of assimilates derived from the fruiting forms themselves is inconclusive. Field experiments were conducted to document the photosynthetic and respiratory activity of cotton leaves, bracts, and capsule walls from anthesis to fruit maturity. Bracts achieved peak photosynthetic rates of 2.1 micromoles per square meter per second compared with 16.5 micromoles per square meter per second for the subtending leaf. However, unlike the subtending leaf, the bracts did not show a dramatic decline in photosynthesis with increased age, nor was their photosynthesis as sensitive as leaves to low light and water-deficit stress. The capsule wall was only a minor site of ¹⁴CO₂ fixation from the ambient atmosphere. Dark respiration by the developing fruit averaged −18.7 micromoles per square meter per second for 6 days after anthesis and declined to −2.7 micromoles per square meter per second after 40 days. Respiratory loss of CO₂ was maximal at −158 micromoles CO₂ per fruit per hour at 20 days anthesis. Diurnal patterns of dark respiration for the fruit were age dependent and closely correlated with stomatal conductance of the capsule wall. Stomata on the capsule wall of young fruit were functional, but lost this capacity with increasing age. Labeled ¹⁴CO₂ injected into the fruit interior was rapidly assimilated by the capsule wall in the light but not in the dark, while fiber and seed together fixed significant amounts of ¹⁴CO₂ in both the light and dark. These data suggest that cotton fruiting forms, although sites of significant respiratory CO₂ loss, do serve a vital role in the recycling of internal CO₂ and therein, function as important sources of assimilate for reproductive development.     

Floral development of Lavatera trimestris and Malva hispanica reveals the nature of the epicalyx in the Malva generic alliance

The epicalyx is a structure below the calyx that is often integrated in floral display. In Malvales, the epicalyx is interpreted to be formed by bracts derived from inflorescence reduction. In this study, we compare the epicalyx and flower development of Lavatera trimestris and Malva hispanica, which are close relatives but show contrasting morphologies. Both species exhibit cymose branching, stipulate subtending leaves, a short plastochron between the appearance of the alternating epicalyx and calyx, a centrifugally developing androecium and a multicarpellar gynoecium. The predominantly trimerous structure and leafy morphology of the epicalyx suggest its origin from a former subtending leaf with leaf‐like stipules. The bilobed epicalyx in M. hispanica represents a loss of the adaxial epicalyx lobe rather than modified bracts. In Malvoideae, the bracts and bracteoles in the flowering branches can be completely absent and are variable in position and number when present. Individual bracts and bracteoles could correspond to further reductions of former subtending leaves instead of precursors of the epicalyx. Although the centrifugal androecium behaves as a branched‐like structure, it is a dynamic complex floral whorl with extended growth capacity. The umbrella in L. trimestris is a swollen part of the style without a well‐understood role in floral or fruit morphology.     

BIFURCATION OF THE STEM APEX IN ASCLEPIAS SYRIACA

In Asclepias syriaca the overall inflorescence is an anthoclad in which the peduncles are non-axillary, each occurring about 60° away from the axil of a leaf. Ontogenetically, a peduncle is initiated when the stem apex expands laterally and bifurcates into separate apices, neither of which is subtended by any type of organ. One of the two apices continues as the functional apex of the stem (bifurcating again at each subsequent node), and the other functions as the apex of the peduncle. The peduncle first produces a bract and, then, a pedicel in the axil of the bract. Subsequent pedicels are each axillary to separate bracts. The pedicels, therefore, can be interpreted as ordinary lateral branches. However, because the bifurcations of the stem apex are not associated with subtending organs, the branching of the stem does not conform to expected monopodial or sympodial systems in the angiosperms. This suggests the possibility that each bifurcation of the stem apex is a true dichotomy. The anthoclad axis, thus, is a series of dichotomies. Although such a series may have been phylogenetically derived from a monopodial or sympodial ancestor, it is also possible that it may have been retained from a primitive, dichotomizing inflorescence.     

ANATOMY AND ASPECTS OF DEVELOPMENT OF THE STAMINATE INFLORESCENCES AND FLORETS OF SEVEN SPECIES OF ALLOCASUARINA (CASUARINACEAE)

The morphology, ontogeny, and vascular anatomy of the staminate inflorescences and florets of seven species of Allocasuarina are described. The generally terminal but open-ended inflorescences occur on monoecious or staminate dioecious trees and consist of whorls of bracts, each subtending a sessile axillary floret. Each floret consists of one terminal stamen with a bilobed, tetrasporangiate anther enclosed typically by cuculliform appendages, commonly considered bracteoles, an inner median pair and an outer lateral pair. The mature stamen is exerted, the anther is basifixed and is extrorsely dehiscent. In early development of a male inflorescence very little internodal elongation occurs and enclosing cataphylls appear. The inflorescence apex is a low dome with a uniseriate tunica and a small group of central corpus cells. Bract primordia are initiated by periclinal divisions of C₁ followed by further divisions of the corpus and anticlinal divisions in the tunica. The bracts are epinastic and become gamophyllous except apically by cell divisions in both sides of each primordium. Stomata are restricted to the axis furrows and the abaxial tips of the bracts. The axillary florets arise in acropetal succession initiated by periclinal divisions in C₁ accompanied by anticlinal divisions in the tunica. The lateral floral appendages are also initiated by C₁ followed by anticlinal divisions in the tunica. They become adnate basally later with the subtending bract. The median sterile appendages are initiated in a manner similar to the initiation of the outer appendages. The stamen is initiated by divisions in the outer layers of the corpus and in the tunica, and then develops first by apical growth followed by intercalary growth. The vascular system of the inflorescence is identical to that of the vegetative stem. Each floret is supplied by a single bundle that has its source in a branch from each of the two traces supplying a bract. Six bundles arise from the floral bundle; four of these terminate in the base of the stamen and two form an amphicribal bundle that supplies the anther. Pollen is binucleate, 3- to 7-porate. The exine is tegillate.     

Initiation of axillary and floral meristems in Arabidopsis

Shoot development is reiterative: shoot apical meristems (SAMs) give rise to branches made of repeating leaf and stem units with new SAMs in turn formed in the axils of the leaves. Thus, new axes of growth are established on preexisting axes. Here we describe the formation of axillary meristems and floral meristems in Arabidopsis by monitoring the expression of the SHOOT MERISTEMLESS and AINTEGUMENTA genes. Expression of these genes is associated with SAMs and organ primordia, respectively. Four stages of axillary meristem development and previously undefined substages of floral meristem development are described. We find parallels between the development of axillary meristems and the development of floral meristems. Although Arabidopsis flowers develop in the apparent absence of a subtending leaf, the expression patterns of AINTEGUMENTA and SHOOT MERISTEMLESS RNAs during flower development suggest the presence of a highly reduced, "cryptic" leaf subtending the flower in Arabidopsis. We hypothesize that the STM-negative region that develops on the flanks of the inflorescence meristem is a bract primordium and that the floral meristem proper develops in the "axil" of this bract primordium. The bract primordium, although initially specified, becomes repressed in its growth.     

Transport of Radioactivity in Relation to Bracts in the Cotton Plant

[¹⁴C]sucrose was applied to the leaf subtending 14-d-old boll(cotton fruit) with and without bracts, and to bracts alonein var. Suvin (Gossypium barbadense L.). The removal of bractssubstantially enhanced the transport of radioactivity from theleaf to the carpels and seed cotton. When bracts alone weretreated, still larger quantities of radioactivity were incorporatedin the carpels and seed cotton. The bracts appeared to nourishthe developing boll through their own photosynthate and regulatethe transport of assimilate from the leaf. Radioactivity, transport, bracts, boll, carpels, seed cotton     

DEVELOPMENTAL STUDIES IN PTYCHOSPERMA (PALMAE). I. THE INFLORESCENCE AND THE FLOWER CLUSTER

This paper describes inflorescence structure, including organogenesis of the panicle and flower clusters and vasculature of flowering branches, for two species of Ptychosperma, a genus of arecoid palms. The inflorescence is an infrafoliar panicle with up to four orders of branches in a spirodistichous arrangement conforming to an irregular one-half phyllotaxy. The primordium of the inflorescence is crescentic and the apex has two tunica layers, a group of central cells, and a rib meristem. The distal flower-bearing parts or rachillae of all branches develop acropetally early in ontogeny and are vertically oriented in the bud. Although these rachillae terminate branches of different sizes and orders, they are similar in size and in number of flower clusters produced. Internodes and lower parts of branches develop later. Bracts of four types are produced: a prophyll and empty peduncular bract, bracts which subtend lateral branches, bracts subtending triads, and floral bracteoles. The prophyll and peduncular bracts are tubular and completely closed around all branches until about three months before the flowers reach anthesis. Bracts subtending lateral branches and those that subtend triads enlarge by small amounts of apical, adaxial, and marginal growth to cover subtended apices during early ontogeny, but are small to absent at maturity. Flower clusters are triads of two lateral staminate and a central pistillate flower. Organogenesis indicates that the triad is a sympodial unit. Flowers develop successively, each floral apex bearing a bracteole that subtends the next flower. The vasculature of the inflorescence may be divided into two systems. Bundles of the main axis extend acropetally into the vertically oriented branches as they are initiated and form a central cylinder of larger bundles in each branch. Flower clusters are supplied by a peripheral system of smaller bundles that develop later in relation to the developing floral organs. Bundles of the peripheral system branch frequently, but branching levels are irregular. The irregular branching of peripheral bundles appears related to the phyllotaxy of the flower clusters and the random right or left position of the first flower of the triad. The level of branching of a bundle may depend on the position of a floral primordium with respect to an existing procambial strand. Three (-4) bundles supply each staminate flower and six (-10) the pistillate flower. The histologically specialized inflorescence has stomata and contains abundant starch. Tannins and raphides, spherical silica bodies, and various forms of sclerenchyma appear in sequence and apparently provide support and protection during the long exposure of the branches.     

Boron Deficiency Induced Changes in Translocation of 14CO2-Photosynthate Into Primary Metabolites in Relation to Essential Oil and Curcumin Accumulation in Turmeric (Curcuma longa L.)

Changes in leaf growth, net photosynthetic rate (P _N), incorporation pattern of photosynthetically fixed ¹⁴CO₂ in leaves 1–4 from top, roots, and rhizome, and in essential oil and curcumin contents were studied in turmeric plants grown in nutrient solution at boron (B) concentrations of 0 and 0.5 g m^-3. B deficiency resulted in decrease in leaf area, fresh and dry mass, chlorophyll (Chl) content, and P _N and total ¹⁴CO₂ incorporated at all leaf positions, the maximum effect being in young growing leaves. The incorporation of ¹⁴CO₂ declined with leaf position being maximal in the youngest leaf. B deficiency resulted in reduced accumulation of sugars, amino acids, and organic acids at all leaf positions. Translocation of the metabolites towards rhizome and roots decreased. In rhizome, the amount of amino acids increased but content of organic acids did not show any change, whereas in roots there was decrease in contents of these metabolites as a result of B deficiency. Photoassimilate partitioning to essential oil in leaf and to curcumin in rhizome decreased. Although the curcumin content of rhizome increased due to B deficiency, the overall rhizome yield and curcumin yield decreased. The influence of B deficiency on leaf area, fresh and dry masses, CO₂ exchange rate, oil content, and rhizome and curcumin yields can be ascribed to reduced photosynthate formation and translocation.     

Rootless Aquatic Plant Aldrovanda Vesiculosa: Physiological Polarity, Mineral Nutrition, and Importance of Carnivory

Various ecophysiological investigations are presented in Aldrovanda vesiculosa, a rootless aquatic carnivorous plant. A distinct polarity of N, P, and Ca tissue content per dry mass (DM) unit was found along Aldrovanda shoots. Due to effective re-utilization, relatively small proportions of N (10 – 13 %) and P (33 – 43 %) are probably lost with senescent leaf whorls, while there is complete loss of all Ca, K, and Mg. The total content of starch and free sugars was 26 – 47 % DM along adult shoots, with the maximum in the 7^th – 10^th whorls. About 30 % of the total maximum sugar content was probably lost with dead whorls. The plant was found to take up 5 – 7 times more NH₄ ⁺ to NO₃ ^– from a mineral medium. Under nearly-natural conditions in an outdoor cultivation container, catching of prey led to significantly more rapid growth than in unfed plants. DM of the fed controls was 48 % higher than in the unfed plants. The controls produced 0.69 branches per plant, while the unfed plants did not produced any. However, the N and P content per DM unit increased by 6 – 25 % in the apices and the first 6 whorls in the unfed variant, as compared to the fed controls. It may be suggested that carnivory is very important for Aldrovanda.     

Leaf characteristics and gas exchange of the mangrove <Emphasis Type="Italic">Laguncularia racemosa</Emphasis> as affected by salinity

Under constant salinity we analysed the leaf characteristics of Laguncularia racemosa (L.) Gaertn. in combination with gas exchange and carbon isotopic composition to estimate leaf water-use efficiency (WUE) and potential nitrogen-use efficiency (NUE). NaCl was not added to the control plants and the others were maintained at salinities of 15 and 30 ‰ (S₀, S₁₅, and S₃₀, respectively). Leaf succulence, sodium (Na), nitrogen (N), and chlorophyll (Chl) contents increased under salinity. Salinity had a negative impact on net photosynthetic rate (P _N) and stomatal conductance (g _s) at high and moderated irradiances. Potential NUE declined significantly (p<0.05) with salinity by 37 and 58 % at S₁₅ and S₃₀, respectively, compared to S₀ plants. Conversely, compared to S₀ plants, P _N/g _s increased under saline conditions by 12 % (S₁₅) and 50 % (S₃₀). Thus, WUE inferred from P _N/g _s was consistent with salinity improved short-term WUE. Long-term leaf WUE was also enhanced by salinity as suggested by significantly increased leaf δ¹³C with salinity. Improved WUE under salinity explains the eco-physiological success of mangrove species under increasing salinity. Conversely, decline in NUE may pose a problem for L. racemosa under hyper-saline environments regardless of N availability.     

Activity of 1-Aminocyclopropane Carboxylic Acid Synthase in Two Mustard (Brassica juncea L.) Cultivars Differing in Photosynthetic Capacity

The pattern of activity of 1-aminocyclopropane carboxylic acid synthase (ACS) was similar to photosynthetic and growth traits observed at 30, 45, and 60 d after sowing in mustard (Brassica juncea L.) cultivars Varuna and RH 30 differing in photosynthetic capacity. Higher activity of ACS and therefore ethylene release in Varuna than RH 30 increased stomatal conductance, intercellular CO₂ concentration, carboxylation rate (carbonic anhydrase and intrinsic water use efficiency), and thus net photosynthetic rate (P _N) and leaf and plant dry masses (DM) at all sampling times. Moreover, Varuna also had larger leaf area which contributed to higher P _N and DM. A positive correlation between ACS activity and P _N and leaf area was found in both the cultivars. Thus ACS activity may affect P _N through ethylene-induced changes on foliar gas exchange and leaf growth.     

Elevated CO2 concentration enhances the role of the ear to the flag leaf in determining grain yield of wheat

Net photosynthetic rate (P _N) of ear and flag leaf during grain filling stage and grain yield of plants with non-darkened or darkened flag leaf or darkened ear were examined in two different CO₂ concentrations: ambient (AC) and AC+200 μmol mol⁻¹ (EC). Ear showed much higher enhancement (56 %) of P _N than flag leaf (23 %) under EC. Moreover, CO₂ enrichment shortened the photosynthetic duration of flag leaf relative to ear. In this way the ratio of ear to flag leaf contribution to grain yield increased from 1.18 (AC) to 1.39 (EC).     

Plant Growth Based on Interrelation between Carbon and Nitrogen Translocation from Leaves

In individual leaves, the photon-saturated photosynthetic activity (P _sat, expressed on a dry mass basis) was closely related to the nitrogen content (Nc) as follows: P _sat = Cf Nc + P _sat0, where Cf and P _sat0 are constants. On a whole plant basis, the relative growth rate (RGR) was closely related to Nc in canopy leaf as follows: RGR = DMf Nc + RGR₀, where DMf and RGR₀ are constants. However, the coefficients Cf and DMf were markedly different among plant species. To explain these differences, it is suggested that carbon assimilation (or dry matter production) is controlled by both the Nc in a leaf (or leaves) and by the net N translocation from leaves. This is supported by the finding that P _sat is related to the rate of ³⁵S-methionine translocation from leaves. We propose another estimation method for the net N translocation rate (NFR) from leaves: Nc, after full leafing, is expressed as a function of time: Nc = (Nc₀ – Ncd) exp(–Nft) + Ncd, where Nf is a coefficient, t is the number of days after leaf emergence, Nc₀ is the initial value of Nc, and Ncd is the Nc of the dead leaf. The NFR is then calculated as NFR = Nc/t = –Nf (Nc – Ncd). Thus Nf is the coefficient for the NFR per unit Nc. NFR is a good indicator of net N translocation from leaves because NFR is closely related to the rate of ³⁵S-methionine translocation from leaves. Since P _sat is related to the ¹⁴C-photosynthate translocation rate, Cf (or DMf) corresponds to the coefficient of saccharide translocation rate per unit amount of Nc. Cf (or DMf) is closely related to the Nf of individual leaves (or the Nf of canopy leaf). This indicates that C assimilation and C translocation from leaves are related to Nc and N translocation from leaves (net translocation of N). Cf and Nf are negatively correlated with leaf longevity, which is important because a high or low CO₂ assimilation rate in leaves is accompanied by a correspondingly high or low N translocation in leaf, and the degree of N translocation in leaves decreases or increases leaf longevity. Thus, since a relatively high P _sat (or RGR) is accompanied by a rapid Nc decrease in leaves, it is difficult to maintain a high P _sat (or RGR) for a sustained time period.     

THE STRUCTURE OF THE ACERVULUS,THE FLOWER CLUSTER OF CHAMAEDOREOID PALMS

Developmental evidence shows that the acervulus, a distinctive flower cluster found only in the chamaedoreoid group of palms, is a form of cincinnus. In Hyophorbe indica Gaertner, the unit consists of a row of sessile flowers, the upper 3–4, staminate and the basal flower, pistillate. During initiation, each new flower originates from divisions in the T₂ and underlying layers of the lower right or left flank of the apex of the preceding flower. A bract subtending the first flower is evident in early stages, is displaced basipetally as the flowers are formed, but is obscured when flowers are mature. No other bracts are associated with the unit. One to two outer bundles of the vascular cylinder of the rachilla develop first to the uppermost flower. Subsequently, bundles to other flowers arise as lower branches of the first bundle and from other, often small outer bundles of the rachilla that become floral traces or produce one or more branches to a flower. Many of the bundles supplying the flowers bend sharply downward in the cortex of the rachilla, apparently reflecting the basipetal sequence of floral inception.