Carbon Dioxide Exchange of Spring-wheat in Relation to Age and Photon-flux Density at Different Growth Temperatures

CO₂-exchange rates (CER) of the sixth and the flag leaves oftwo spring-wheat varieties, Kolibri and Famos, were comparedusing an open-circuit infrared gas analysing system. Measurementswere repeated every two weeks starting when leaf blades werefully expanded. Single plants were grown in a controlled environmenthaving a photopuiod of 15 h and a day/night temperature of 24/19°C(H), 18/13 °C (M), and 12/7 °C (L) respectively untilapprox. 2 weeks after anthesis and at 18/13 °C until maturity.The photosynthetic photon-flux density (PPFD) at the top ofthe plants was 500 µE m^–2 sec^–1. During themeasurements PPFD was gradually reduced from 2000 to 0 µEm^–2 sec^–1 whereas the temperature was maintainedat the respctive growth-temperatures during the light period.The CER of the sixth leaf declined fairly similarly for bothvarieties, except for Kolibri where a faster decline was observedduring the first two weeks after full leaf expansion. The CERof the flag leaf declined more slowly than that of the sixthleaf. With the flag leaf of Famos, the decline was nearly linear,whereas with Kolibri it was very slow during the first few weeksbut rapid as the leaves further senesced. This pattern becamemore pronounced as the growth temperature decreased. The declinein relation to leaf age was much smaller at low PPFD than athigh PPFD during the same period. At full leaf expansion Kolibrireached higher maximum CER than Famos except at H. As the PPFDwas reduced the difference became smaller and at very low PPFDsuch as 50 µE m^–2 sec^–1 was reversed for thesixth leaf. Under optimum growth conditions maximum values ofCER were greater than 50mg CO₂ dm^–2h^–1 and PPFDfor light saturation was close to 2000 µE m^–2 sec^–1.A comparison between the actual CER and a fitted curve widelyused, PN=(a+b/l)^–1–DR, showed that the goodnessof fit strongly depends on cultivar, treatment and leaf ageas well as on the number and the level of PPFD from which datafor calculations are taken. Triticum aestivum, L., wheat, photosynthesis, photon-flux density, light response, carbon, dioxide exchange     

Boron supply into wheat (Triticum aestivum L. cv. Wilgoyne) ears whilst still enclosed within leaf sheaths

The present study investigates whether there is significant remobilization of (10)B previously loaded in the flag and penultimate leaves into the young, actively growing ear enclosed within the sheaths of flag and penultimate leaves. It also explores whether B transport into the enclosed ear declines when air humidity in the shoot canopy increases. After 5 d (10)B labelling during the period from early to full emergence of the flag leaf, the plants were transferred into nutrient solutions containing either 10 microM (11)B or no added B for 3 d. Regardless of the subsequent B supply levels to the roots, (10)B contents in the ear continued to increase by up to 5-fold 3 d after the end of (10)B supply in the nutrient solution. During these 3 d, the ear experienced a rapid increase in biomass. However, the majority of B in the ear during the 3 d treatment period was from the newly acquired (11)B from root uptake, rather than retranslocation of (10)B previously deposited in the leaves. By comparing the relative distribution of (10)B, Rb (xylem-to-phloem transfer marker) and Sr (xylem-marker) in the ear and the flag leaf, the distribution of (10)B resembled that of Rb more than Sr. Canopy cover treatment greatly suppressed leaf transpiration and decreased the amount of newly acquired (10)B in the flag leaf and the ear, but not in the upper stem segments. The results suggest that whilst the young ear was still fully enclosed within the leaf sheaths without any significant transpiration activity, B transport into the ear is predominantly dependent on the long-distance B transport in the xylem driven by leaf transpiration and, therefore, on concurrent B uptake from the roots.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE L.)

During the first year of growth of timothy, variations in nitrogen supply affected the number of tillers and leaves, leaf area and dry weight, but had little influence on relative growth and net assimilation rates, except at very low levels of nitrogen. Differentiation and ear emergence occurred slightly earlier at high nitrogen and ear length was increased; there was also an increase in the proportion of fertile tillers among the total number produced after the first few weeks of tillering. The effect of nitrogen supply on the number, dry weight and ear length of early primary tillers was small but became larger in later tillers. Irrespective of the level of nitrogen supply, tillers had particularly high net assimilation rates during the first few weeks following their appearance.     

Modification of flag leaf senescence and yield characters in barley (Hordeum vulgare L.) by gibberellic acid and kinetin

Barley plants (Hordeum vulgare L.) received foliar applications of 10^?4 M gibberellic acid (GA₃) and Kinetin (KN) individually and in combination at one or more of three growth stages: flag leaf appearance (I), ear emergence (II), and the first stage of senescence initiation in the flag leaf (III). Both plant growth regulators (PGR) hastened onset of senescence when sprayed at Stage I and/or Stage II. Treatment at Stage III, either alone or in combination with treatments at the other stages, tended to postpone senescence. Yield components also showed stage-dependent response: Stage I treatment increased the formation of total and bearing tillers, and Stage III treatment improved grain number and weight. However, while GA₃ proved more effective than KN, the two together acted antagonistically.     

Ontogenetic variations on absorption and utilization of phosphorus in common bean cultivars under biological nitrogen fixation

Salinity is a major yield-reducing factor in coastal and arid, irrigated rice production systems. Salt tolerance is a major breeding objective. Three rice cultivars with different levels of salt tolerance were studied in the field for growth, sodium uptake, leaf chlorophyll content, specific leaf area (SLA), sodium concentration and leaf CO₂ exchange rates (CER) at photosynthetic active radiation (PAR)-saturation. Plants were grown in Ndiaye, Senegal, at a research station of the West Africa Rice Development Association (WARDA), during the hot dry season (HDS) and the wet season (WS) 1994 under irrigation with fresh or saline water (flood water electrical conductivity = 3.5 mS cm^-1). Relative leaf chlorophyll content (SPAD method) and root, stem, leaf blade and panicle dry weight were measured at weekly intervals throughout both seasons. Specific leaf area was measured on eight dates, and CER and leaf sodium content were measured at mid-season on the first (topmost) and second leaf. Salinity reduced yields to nearly zero and dry-matter accumulation by 90% for the susceptible cultivar in the HDS, but increased leaf chlorophyll content and CER at PAR- saturation. The increase in CER, which was also observed in the other cultivars and seasons, was explained by a combination of two hypotheses: leaf chlorophyll content was limited by the available N resources in controls, but not in salt-stressed plants; and the sodium concentrations were not high enough to cause early leaf senescence and chlorophyll degradation. The growth reductions were attributed to loss of assimilates (mechanisms unknown) that must have occurred after export from the sites of assimilation. The apparent, recurrent losses of assimilates, which were between 8% and 49% according to simulation with the crop model for potential yields in irrigated rice, ORYZA S, might be partly due to root decomposition and exudation. Possibly more importantly, energy-consuming processes, such as osmoregulation, interception of sodium and potassium from the transpiration stream in leaf sheaths and their subsequent storage, drained the assimilate supply. This revised version was published online in August 2006 with corrections to the Cover Date.     

Modification of flag leaf senescence and yield characters in barley (Hordeum vulgare L.) by gibberellic acid and kinetin

Barley plants (Hordeum vulgare L.) received foliar applications of 10^–4 M gibberellic acid (GA₃) and Kinetin (KN) individually and in combination at one or more of three growth stages: flag leaf appearance (I), ear emergence (II), and the first stage of senescence initiation in the flag leaf (III). Both plant growth regulators (PGR) hastened onset of senescence when sprayed at Stage I and/or Stage II. Treatment at Stage III, either alone or in combination with treatments at the other stages, tended to postpone senescence. Yield components also showed stage-dependent response: Stage I treatment increased the formation of total and bearing tillers, and Stage III treatment improved grain number and weight. However, while GA₃ proved more effective than KN, the two together acted antagonistically.     

Effect of drought on ear and flag leaf photosynthesis of two wheat cultivars differing in drought resistance

We investigated net photosynthetic rate (P_N) of ear and two uppermost (flag and penultimate) leaves of wheat cultivars Hongmangmai (drought resistant) and Haruhikari (drought sensitive) during post-anthesis under irrigated and non-irrigated field conditions. The P_Nof ear and flag leaf were significantly higher and less affected by drought in Hongmangmai than in Haruhikari. The rate of reduction in stomatal conductance (g_s) was similar for the two cultivars, but intercellular CO₂concentration (C_i) in the flag leaf of Hongmangmai was lower than that of Haruhikari in non-irrigated treatment. No differences were observed in leaf water potential (₁) and osmotic adjustment of the flag leaf of the cultivars. These results imply that differences in photosynthetic inhibition on the flag leaf at low leaf ₁between the cultivars were primarily due to non-stomatal effects. Hence the main physiological factor associated with yield stability of Hongmangmai under drought stress may be attributed to the capacity for chloroplast activity in the flag leaf, which apparently allows sustained P_Nof flag leaf during grain filling under drought stress. The higher P_Nof ear in Hongmangmai under drought could also be related to its drought resistance.This revised version was published online in March 2005 with corrections to the page numbers.     

Transpiration in Barley Lines with Differing Stomatal Frequencies

Leaf conductances and transpiration rates from potted plantswere studied on two pairs of barley lines selected for highor low stomatal frequency on the flag leaf. Although there werelarge differences in stomatal frequency on the flag leaves,there was no evidence that the low frequency lines had the lowerconductances at equal leaf water potentials. This may have beendue to the changes in the size of the stomata which compensatedfor differences in stomatal frequency. Although there were no differences in stomatal conductance (expressedon a unit leaf area basis) the rate of water use per plant wasup to 50% faster for the low frcquency lines than for the high,particularly after emergence of the sixth leaf. This was causedby a larger green leaf area at this stage which was in turndue to larger individual leaves, more tillers, and a slowersenescence of the older leaves. These observations are discussed in relation to the possibilityof selecting for stomatal characteristics as a means of breedingvarieties able to tolerate drought.     

Photosynthetic Characteristics of Non-Leaf Organs of Winter Wheat Cultivars Differing in Ear Type and their Relationship with Grain Mass Per Ear

Chlorophyll content, photosystem 2 functioning (F_v/F_m, F_v/F₀), activity of ribulose-1,5-bisphosphate carboxylase/oxygenase, and net photosynthetic rates (P _N) of flag leaf blade, sheath, peduncle, and ear organs were assessed in large-ear type (Pin 7) and small-ear type (ND93) wheat cultivars. Some differences were found in photosynthetic properties between different green plant parts, the values of all studied parameters in ear parts being higher in Pin7 than in ND93. Furthermore, ear surface areas and ear P _N in 26 wheat genotypes measured at anthesis showed highly significant positive correlation with grain mass per ear. Hence a greater capability of ear photosynthesis may result in a greater grain yield in large-ear type cultivars.     

Soil salinity effects on transpiration and net photosynthetic rates,stomatal conductance and Na+ and Cl− contents in durum wheat

Leaf gas exchange, plant growth and leaf ion content were measured in wheat (Triticum durum L. cv. HD 4502) exposed to steady- state salinities (1.6, 12.0 and 16.0 dS nr⁻¹) for 8 weeks. Salinity reduced leaf area and number of tillers, and increased Na⁺ and Cl⁻ concentrations in leaves. Leaf- to- leaf gradients of these ions were observed. The oldest leaf contained 6 to 8 times more Na⁺ and Cl⁻ than the flag leaf. Net photosynthetic rate (P_N), transpiration rate (E) and stomatal conductance (g_S) were the highest in flag leaf, declined in the middle and fully expanded leaves, and were minimum in the oldest leaves. These processes were reduced by salinity with similar leaf- to- leaf gradients. Intercellular CO₂ concentrations in the older leaves were higher than in the flag leaf in non-saline plants, and increased similarly with salinity. Leaf age was the major factor in reducing P_N, and senescence processes were promoted by salinity.     

RuBP carboxylase activity in wheat following ear emergence and evaluation of the role of ear

Ribulose bisphosphate carboxylase activity and chlorophyll content were measured in the leaves and reproductive parts of two wheat cultivars after ear emergence. The chlorophyll content of the flag leaf was mostly higher than that of the awns and glumes. Awns had the highest chlorophyll content among reproductive parts. Light transmission to the lower leaves was higher in the dwarf cultivar Moti than in the medium tall cultivar Kalyansona. RuBP carboxylase activity in Kalyansona leaves was higher than in Moti leaves. In postanthesis stages there was no difference in RuBP carboxylase activity in the flag leaf and lower leaves between cultivars. Awns had the maximum activity of RuBP carboxylase followed by glumes and grains among ear parts. The relative capacity for photosynthesis in the ear parts was several times higher than in the flag leaf on a unit chlorophyll basis. It is suggested that in a crop canopy in the field, the spike(ear) may have a greater importance in grain development than has been previously estimated.     

Stomatal control of gas exchange in barley awns

The gas exchange of barley ears and awns was measured in the field using a gas analysis system and a diffusion porometer. Awn stomatal resistance decreased with increasing irradiance but to a smaller extent than leaf stomatal resistance. Measurements on ears immediately before and after successively removing awns showed that awn transpiration and photosynthesis were proportional to awn area and that awns accounted for 73% of transpiration by the ear. Although the maximum rates of photosynthesis of which awns were capable declined with age, awns accounted for 80–115% of the net CO₂ uptake of complete ears because the ears-less-awns could respire more CO₂ than they absorbed. Ear photosynthesis accounted for 52% of the weekly increment in ear dry weight after ear emergence, but 5 weeks later photosynthesis by the ear balanced respiration. Overall photosynthesis by the ear accounted for 35 % of its final weight. Differences in the light response curves of leaves and ears can be fully accounted for by the different relationships between stomatal resistance and irradiance of the two organs.     

Carbon dioxide exchange in relation to sink demand in wheat

Summary In this paper, experiments are described which examine the effect of requirement for assimilates by the ear on the rate of net photosynthesis in leaves of wheat (Triticum aestivum L.). Different levels of requirement were achieved by various levels of sterilization of florets just before anthesis, which resulted in a range of grain numbers per ear, and by inhibiting photosynthesis of the intact ear by 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU). Only the ear and two uppermost leaves of the main shoot were considered, all the lower leaves and tiller leaves being excised when the experimental treatments were imposed. In two experiments, tiller regrowth was permitted during the experimental period, while in a third, new tillers were defoliated regularly.The response of leaf photosynthesis to the level of assimilate requirement by the ear was influenced by the treatment of the vegetative tillers. Thus, the net photosynthesis rate of the flag leaf was decreased by a reduction in grain number, or increased by inhibition of photosynthesis in the ear, only when the vegetative tillers were kept defoliated; when these tillers were allowed to re-grow normally, there was no influence of ear treatment on leaf photosynthesis. Temporal changes in leaf photosynthesis were consistent with this response pattern, i.e., when tillers were defoliated, the initial high rates of photosynthesis persisted for much longer.In the experiment where photosynthesis was influenced by the requirement for assimilate in the ear, the variation occurred through change in stomatal conductance on the abaxial surface of the leaf. This surface has a lesser conductance to CO₂ exchange than the adaxial surface. The implication of this finding to rapid methods of plant screening is discussed.     

Effect of drought on ear and flag leaf photosynthesis of two wheat cultivars differing in drought resistance

We investigated net photosynthetic rate (P_N) of ear and two uppermost (flag and penultimate) leaves of wheat cultivars Hongmangmai (drought resistant) and Haruhikari (drought sensitive) during post-anthesis under irrigated and non-irrigated field conditions. The P_Nof ear and flag leaf were significantly higher and less affected by drought in Hongmangmai than in Haruhikari. The rate of reduction in stomatal conductance (g_s) was similar for the two cultivars, but intercellular CO₂concentration (C_i) in the flag leaf of Hongmangmai was lower than that of Haruhikari in non-irrigated treatment. No differences were observed in leaf water potential (ψ₁) and osmotic adjustment of the flag leaf of the cultivars. These results imply that differences in photosynthetic inhibition on the flag leaf at low leaf ψ₁between the cultivars were primarily due to non-stomatal effects. Hence the main physiological factor associated with yield stability of Hongmangmai under drought stress may be attributed to the capacity for chloroplast activity in the flag leaf, which apparently allows sustained P_Nof flag leaf during grain filling under drought stress. The higher P_Nof ear in Hongmangmai under drought could also be related to its drought resistance.     

Effect of assimilate utilization on photosynthetic rate in wheat

Summary Two weeks after anthesis, when the grain is filling rapidly, the rate of photosynthesis by flag leaves of wheat cv. Gabo was between 20 and 30 mg CO₂ dm^-2 leaf surface hour^-1 under the conditions used. About 45% of flag-leaf assimilates were translocated to the ear, and only about 12% to the roots and young shoots.On removing the ear, net photosynthesis by the flag leaves was reduced by about 50% within 3–15 hours, and there was a marked reduction in the outflow of ¹⁴C-labelled assimilates from the flag leaves.Subsequent darkening of all other leaves on plants without ears led to recovery of flag-leaf photosynthesis, as measured by gas analysis and ¹⁴CO₂ fixation, and to increased translocation of assimilates to the roots and young shoots. Minor changes in the rates of dark respiration accompanied these major, reversible changes in photosynthetic rate.After more than a week in continuous, high-intensity light, the rate of photosynthesis by flag leaves of intact plants had fallen considerably, but could be restored again by a period in darkness, or by inhibiting photosynthesis in the ears by spraying them with DCMU. The inhibition of ear photosynthesis increased translocation of labelled assimilates from the flag leaf to the ears, without affecting leaf sugar levels.The application of TIBA to the culm below the ear inhibited auxin movement throught the culm, but had no influence on flag-leaf photosynthesis.These results suggest that, at least in this system, photosynthesis by the flag leaf is regulated directly by the demand for assimilates from the flag leaf and not indirectly through action in the leaf of auxins produced by the sink organs.     

Effects of Water Stress on Photosynthesis and Carbon Partitioning in Soybean (Glycine max [L.] Merr.) Plants Grown in the Field at Different CO(2) Levels

The effects of water stress and CO₂ enrichment on photosynthesis, assimilate export, and sucrose-P synthase activity were examined in field grown soybean plants. In general, leaves of plants grown in CO₂-enriched atmospheres (300 microliters per liter above unenriched control, which was 349 ± 12 microliters per liter between 0500 and 1900 hours EST over the entire season) had higher carbon exchange rates (CER) compared to plants grown at ambient CO₂, but similar rates of export and similar activities of sucrose-P synthase. On most sample dates, essentially all of the extra carbon fixed as a result of CO₂ enrichment was partitioned into starch. CO₂-enriched plants had lower transpiration rates and therefore had a higher water use efficiency (milligrams CO₂ fixed per gram H₂O transpired) per unit leaf area compared to nonenriched plants. Water stress reduced CER in nonenriched plants to a greater extent than in CO₂-enriched plants. As CER declined, stomatal resistance increased, but this was not the primary cause of the decrease in assimilation because internal CO₂ concentration remained relatively constant. Export of assimilates was less affected by water stress than was CER. When CERs were low as a result of the imposed stress, export was supported by mobilization of reserves (mainly starch). Export rate and leaf sucrose concentration were related in a curvilinear manner. When sucrose concentration was above about 12 milligrams per square decimeter, obtained with nonstressed plants at high CO₂, there was no significant increase in export rate. Assimilate export rate was also correlated positively with SPS activity and the quantitative relationship varied with CER. Thus, export rate was a function of both CER and carbon partitioning.     

Boron accumulation and partitioning in wheat cultivars with contrasting tolerance to boron deficiency

Two pot experiments at the Plant Environment Laboratory (PEL), Reading, UK investigated sterility, boron (B) accumulation and B partitioning of wheat cultivars grown with limited B in the growing medium. The first experiment evaluated nine cultivars of spring wheat with diverse field responses to low available soil B, supplied with or without 20 μM B. A second experiment examined the response of a susceptible (SW-41) and a tolerant (Fang-60) cultivar to B-deficiency. These cultivars were supplied with either 20 μM B from sowing to flag leaf emergence and no added B thereafter, or 20 μM B from sowing to maturity. When B was not supplied in the nutrient solution, the number of grains ranged from 4 per ear (cv. BL-1135) to 32 per ear (cv. BL-1249) and sterility of competent florets ranged from 39% to 93%. Boron concentration in the flag leaf at anthesis did not differ greatly when the growing medium contained limited B, but differences between cultivars were evident when B was unlimited. Tolerance of B-deficiency was not related to the B concentration in the flag leaf. Some cultivars produced viable pollen and set grains while others failed to do so at similar B concentrations in the flag leaf. The two contrasting cultivars did not differ much in their pattern of B partitioning when B supply was restricted from flag leaf emergence onwards. Similarly, little evidence was found that the tolerant cultivars translocated B from their leaves, roots or stems when the supply in the growing medium was restricted. The proportion of total B partitioned in different organs was the same irrespective of B supply and cultivar. On average, leaves contained 68% of the total B content in the whole plant compared to 16% in the roots, 10% in the ears and only 6% in the stems. Tolerant or susceptible cultivars of wheat could not be distinguished based on the B concentration and B content of the flag leaf. This revised version was published online in June 2006 with corrections to the Cover Date.     

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).     

Correlations among leaf CO2-exchange rates,areas and enzyme activities among soybean cultivars

Soybean (Glycine max (L.) Merr.) genotypes varying in area per nodal unit (usually a trifoliolate) and maturity class were grown in plots at the University of Illinois experimental farm. Leaf CO₂-exchange rates per unit area (CER) were measured under sunlight on intact plants. In addition to previously reported correlations with specific leaf weight and chlorophyll, CER was positively correlated with ribulose bisphosphate carboxylase (RuBPcase) activity, specific activity, and soluble protein, and was negatively correlated with area per leaf unit. The CER: chlorophyll correlation was destroyed by high CER values in 2 chlorophyll-deficient lines. CER values for 27 of the 35 lines tested fell within the range of those for isolines of cultivar Clark varying in leaf characteristics. The CER values were highest for fully expanded leaves during rapid pod fill. These results suggested that photoperiod (maturity) genes and genes for leaf area growth interact with genes controlling photosynthetic CO₂-exchange to produce the major differences in CER values among soybean genotypes.     

Ear photosynthesis, carbon isotope discrimination and the contribution of respiratory CO2 to differences in grain mass in durum wheat

The role of ear photosynthesis in grain filling was studied in a number of durum wheat (Triticum turgidum var durum L.) landraces and varieties from the Middle East, North Africa, and from the collections of ‘Institut National de la Recherche Agronomique’ (INRA, France) and ‘Centro International de Mejora de Maiz y Trigo’ (CIMMYT, Mexico). Plants were grown in the field in a Mediterranean climate. Flag leaves (blade plus sheath) and ears were kept in the dark from 1 week after anthesis to maturity which reduced grain weight by 22.4% and 59.0%, respectively. In a further experiment, the carbon isotope discrimination ratio (Δ) of ear bracts, awns and flag leaves was measured on samples taken at anthesis and on mature kernels. The mean value of Δ for the water soluble fraction of bracts (17.0‰) and awns (17.7‰) were lower than those of leaves (19.5‰) and fairly similar to those of kernels (17.4‰) averaged across all genotypes. Data indicate that most of the photosynthates in the grain come from ear parts and not from flag leaves. In addition, a higher water use efficiency (WUE) of ear parts than of the flag leaf is suggested by their lower Δ values. Gas exchange in ears and flag leaves was measured during grain filling. Averaged over all genotypes, CO₂ diffusive conductance was about five times higher in the flag leaf than in the spike (with distal portions of awns outside the photosynthetic chamber) 2 weeks after anthesis. In absolute terms, the dark respiration rate (R_d) was greater than the net photosynthesis rate (P_n) by a factor of 1.74 in the spike, whereas R_d was much smaller, only 22.1, 65.7 and 24.8% of P_n in blade, sheath and awns, respectively. Data indicate that photosynthesis, and hence the water use efficiency (photosynthesis/transpiration), is greatly underestimated in ears because of the high rates of respiration which diminish the measured rates of net CO₂ exchange. Results of ¹³C discrimination and gas exchange show that genotypes from North Africa have higher WUE than those from the Middle East. The high R_d values of ears as well as their low diffusive conductance suggest that CO₂ from respiration may be used as source of carbon for ear photosynthesis. In the same way, the anatomy of glumes, for example, supports the role of bracts using internal CO₂ as source of photosynthesis. In the first experiment, the Δ in mature grains from culms with darkened ears compared with control culms provided further evidence in support of this hypothesis. Thus, the Δ from kernels of control plants was 0.40 higher than that from ear-darkened plants, probably because of some degree of refixation (recycling) of respired CO₂ in the grains.