Effect of Low Relative Humidity on {delta}13C Value in Two C3 Grasses and in Panicum milioides, a C3-C4 Intermediate Species

When grown under conditions of low relative humidity, the C₃–C₄intermediate Panicum milioides, as well as the C₃ grasses Triticumaestivum and Poa pratense, exhibited ¹³C values which were upto 2–7%o less negative than the ¹³C values of the correspondingplants grown at high relative humidity. At both humidity levels,there was no evidence of a substantial contribution of phosphoenolpyruvatecarboxylase to carbon gain in Panicum milioides     

Intracellular Silica Deposition in Immature Leaves in Three Species of the Gramineae

The formation of solid, discrete deposits of opaline silicawithin the cell lumen of leaf tissues is reported in speciesrepresenting three subfamilies of the Gramineae; the preparationof a silica-minimal nutrient solution is discussed. Opal phytolithnumbers are related to tissue age and to two external silicaconcentrations for tiller leaves over a period of 32 days followingbud initiation. Variations in silica deposition patterns among the individualleaves of a homologous series on the shoot apex are relatedto differential growth-rates. During ontogeny, deposition occursin an ordered sequence of cell types related to the basipetalmaturation gradient within the leaf. Initial deposition wasdetected in silica cells (idioblasts) of imbricated, bud leavesof Sieglingia decumbens, 1 mm long; phytoliths are confinedto these cells in expanding, basal portions of the leaf. Phytolithcounts/sq mm of epidermis vary with the degree of long-cellexpansion. The error is reduced by expressing silicificationas the number of phytoliths/100 silica cells. Post-expansiondeposition was initiated in epidermal long cells of the leafblade tip. The higher silica concentration resulted in a morerapid utilization of available deposition sites and larger phytoliths(P = 0.001). A passive cell influx of silicic acid is discussedin relation to cytodifferentiation.     

Photosynthetic Characteristics of C3-C4 Intermediate Flaveria Species II. Kinetic Properties of Phosphoenolpyruvate Carboxylase from C3, C4 and C3-C4 Intermediate Species

The kinetic properties of phosphoenolpyruvate (PEP) carboxylasehave been studied among several Flaveria species: the C₃ speciesF. cronquistii, the C₃–C₄ species F. pubescens and F.linearis, and the C₄ species F. trinervia. At either pH 7 or8, the maximum activities (in µmol.mg Chl^–1.h^–1)for F. pubescens and linearis (187–513) were intermediateto those of the C₃ species (12–19) and the C₄ species(2,182–2,627). The response curves of velocity versusPEP concentration were hyperbolic for the C₃ and C₃–C₄species at either pH 7 or 8 while they were sigmoidal for theC₄ species at pH 7 and hyperbolic at pH 8. The K_m values forPEP determined from reciprocal plots were lowest in the C₃ species,and of intermediate value in the C₃–C₄ species comparedto the K' values of the C₄ species determined from Hill plotsat either pH 7 or 8. Glucose-6-phosphate (G6P) decreased theK_m values for PEP at both pH 7 and 8 in the C₃ and C₃–C₄species. In the C₄ species, G6P decreased the K' values at pH8 but increased the K' values at pH 7. In all cases, G6P hadits effect by influencing the activity at limiting PEP concentrationswith little or no effect on the maximum activity. At pH 8 andlimiting concentrations of PEP the degree of stimulation ofthe activity by G6P was greatest in the C₄ species, intermediatein F. linearis, a C₃–C₄ species, and lowest in the C₃species. In several respects, the PEP carboxylases of the C₃–C₄Flaveria species have properties intermediate to those of theC₃ and C₄ species. (Received April 30, 1983; Accepted August 22, 1983)     

Discriminating and mapping the C3 and C4 composition of grasslands in the northern Great Plains,USA

There is uncertainty about the extent and distribution of grasslands following the C3 and C4 photosynthetic pathways. Since these grasses have an asynchronous seasonal profile it should be possible to estimate and map the C3–C4 composition of grasslands from multi-temporal remote sensing imagery. This potential was evaluated using 30 weekly composite MERIS MTCI images for South Dakota, USA. Derived relationships between the remotely sensed response and composition of grasslands were significant, with R² ∼ 0.6. It also appears possible to map broad classes of grassland composition, with a three class (high, medium and low C3 cover) classification having an accuracy of 77.8%.     

Diel Acid Fluctuations in C4 Amphibious Grasses

Orcuttieae is a small tribe of C4 grasses endemic to seasonal pools in the southwestern U.S., comprising the basal genus Neostapfia, Tuctoria, and the most derived group, Orcuttia. Growth is initiated underwater, and when pools dry, species undergo a metamorphosis replacing aquatic foliage with terrestrial foliage. O. californica and O. viscida exhibit CAM-like diel fluctuations in acidity in the aquatic foliage. Pulse-chase studies showed that although CO2 was fixed into malic acid in the dark, an overnight chase in the dark revealed that most label was not retained in organic acids, indicating a role other than CAM. Terrestrial foliage exhibited a very different diel fluctuation; acids accumulated during the day, and diminished overnight. Malic acid predominated and was secreted on the surface of the leaf in a manner similar to another arid land species. This terrestrial daytime acid accumulation may not be related to photosynthetic pathway but may play an anti-herbivore function. No acid fluctuations were observed in either N. colusana or T. greenei.     

Silica and Ash in Tissues of Some Plants Growing in the Coastal Area of Mississippi, USA

Ash and silica contents and their depositional patterns in differenttissues of 27 plants growing in the Ocean Springs area of Mississippi(many grow elsewhere), were determined. Silica content of driedplant parts varied from no more than 0 per cent in Borrichiafrutescens (L.) D.C. stems to 18.76 per cent in Arundinariagigantea (Walt.) Muhl. leaves. Ash content varied from 0·73per cent in Cliftonia monophylla (Lam.) Britt. ex Sarg. stemsto 44·02 per cent in Batis maritima L. leaves. Plantssuch as Batis maritima L., Borrichia frutescens (L.) D.C., Salicorniabigelovii Torr. and Salicornia virginica L. which grew in salinemarshes had high ash contents due NaCl in their tissues. Morusrubra L. leaves had a high silica content for a dicotyledonousplant (3·12 per cent). Energy-dispersive X-ray analysisshows that the distribution of the element silicon is clearlyrelated to certain epidermal structures such as ridges, cellwalls, rows of irregular shaped structures lengthwise of theleaf, dumb-bell shaped ones and especially in trichomes. Therewas a high concentration of silica containing trichomes alongthe veins on the underside of Morus rubra L. leaves and thiswould protect them from insects. The outer parts of the inflorescencesof Ctenium aromaticum (Walt.) Wood, Elymus virginicus L., Juncuspolycephalus Michx. and phragmites communis Trin. were highlysilicified. This should give the seed some protection from insects.The sharp particles would be irritating to oesophageal tissuesand might be implicated in such a cancer.     

Effects of Anoxia on Phloem Loading in C3 and C4 Species

Changes in phloem loading rate were inferred from observationsof ¹¹C export from attached leaves of C₃ and C₄ monocots anddicots. Loading decreased under anoxia in C₃ leaves, but notin general in the leaves of either C₄ monocots or dicots inthe light. However, loading rate in the C₄ leaves did reduceif the leaf was also darkened or received no CO₂. We suggestthat insensitivity to anoxia in C₄ leaves is due to oxygen fromphotosynthesis which is retained in the bundle sheath at a concentrationsufficient to energize a phloem loading system. There may alsobe another system which is insensitive to anoxia since the effectsof shade and CO₂ deprivation were not always seen, and loadingwas not completely stopped by these treatments. Key words: Phloem loading, C₃, C₄     

Quantity and Kinetic Properties of Ribulose 1,5-Bisphosphate Carboxylase in C3, C4, and C3-C4 Intermediate Species of Flaveria (Asteraceae)

The kinetic properties of ribulose 1,5-bisphosphate carboxylase(RuBPC) appear to have been modified during evolution of photosynthesisto adjust to changes in substrate availability. C₄ plants areconsidered to have a higher concentration of CO₂ available toRuBPC than C₃plants. In this study, the K_m(CO₂ and catalyticcapacity (k_cat) of RuBPC and the ratio of RuBPC protein to totalsoluble protein from several Flaveria species, including C₃,C₃-C₄ intermediate, and C₄ species, were determined. The C₃and intermediate species had similar K_m(CO₂) values while theC₄ species on average had higher K_m(CO₂) values. The mean ratioof K_cat/K_m for species of each group was similar, supportingthe hypothesis that changes in K_m and K_cat, are linked. Theallocation of total soluble protein to RuBPC was lowest in theC₄ Flaveria species, intermediate in the C₃-C₄ species, andhighest in the C₃ species. The results suggest that during evolutionof C₄ photosynthesis adjustments may occur in the quantity ofRuBPC prior to changes in its kinetic properties. (Received January 4, 1989; Accepted April 11, 1989)     

Gas Exchange and Chlorophyll Fluorescence of C3 and C4 Saltmarsh Species

Spartina maritima (Curtis) Fernald, Spartina densiflora Brong, Arthrocnemum perenne (Miller) Moss, and Arthrocnemum fruticosum (L.) Moq are very frequent halophytes on the coasts of SW Europe. The first two are perennial Gramineae with C₄ metabolism; the last two are perennial Chenopodiaceae with C₃ metabolism. Controlled garden experiments were carried out with the four species to compare their physiological response, i.e., water potential (Ψ), net photosynthetic rate (P_N), transpiration rate (E), stomatal conductance (g_s), intercellular CO₂ concentration (Ci), and chlorophyll fluorescence of photosystem (PS) 2 under saline and non-saline conditions. S. maritima behaves as an osmoconformer species, the other three as osmoregulators. In the four species, P_N, E, and g_s improved following freshwater irrigation. The variations in P_N might be related with biochemical changes (which appear not to affect PS2), but not with significant stomatal fluctuations, which are associated with a lower water use efficiency in the case of Arthrocnemum. The species were segregated into two groups (not depending on their C₃ or C₄ photosynthetic pathway), in relation with the topographic level of this species in natural conditions: the relative responses of P_N in S. maritima and A. perenne were lower than those of S. densiflora and A. fruticosum. The salt-tolerance index supports such segregation. S. densiflora demonstrated the best competitive possibilities against salt-tolerant glycophytes, with its more flexible response in saline or brackish environments, which explains its spreading along the rivers draining into the estuaries of the SW Iberian Peninsula. This revised version was published online in June 2006 with corrections to the Cover Date.     

Comparative Anatomy and Morphology of Leaves between C3 and C4 Species in Panicum

Anatomical and morphological structures of leaf blades werecompared between C₃ and C₄ species in Panicum. Inter-specificvariation of stomatal density, longitudinal vein density andmesophyll thickness was highly correlative either plus or minuswithin respective groups. The two groups could not be distinguishedby a single character, since the variation ranges overlappedeach other. However, the quantitative relations between veindensity and the other two characters differentiated the twogroups well. In C₃, stomatal density seemed to be a primaryfactor for regulating water balance, while in C₄ vein systemwas considered to be important for the regulation. The specieswith intermediate photosynthesis behaved similar to the C₃ species.In the C₃ group, correlative variation was observed betweenleaf width, leaf angle and the three characters mentioned above.Variation of light-receiving area due to the changes of widthand angle of leaf blades was considered to be one of the adaptivestrategies of this group. Increase of light-receiving area wasin connection with the thinning of leaves. On the other hand,in the C₄ correlations between length, width and angle of leaveswere low. Such loose character correlation may be achieved byits efficiency of CO₂ utilization and its well developed veinsystems. Besides, NAD-me type species tended to have relativelylower stomatal and vein densities as compared with the otherdecarboxylation types in this group. Panicum, photosynthesis, C₃, C₄, decarboxylation types, leaf, stomata, vein     

Regulation of Electron Transport in Photosystems I and II in C3, C3-C4, and C4 Species of Panicum in Response to Changing Irradiance and O2 Levels

Regulation of the quantum yields of linear electron transport and photosystem II photochemistry ([phi]II) with changing irradiance and gas-phase O2 concentration was studied in leaf tissue from Panicum bisulcatum (C3), Panicum milioides (C3-C4), and Panicum antidotale (C4) at 200 [mu]bars of CO2 and 25[deg]C using infrared gas analysis and chlorophyll fluorescence yield measurements. When the O2 level was increased from 14 to 213 mbars at high irradiance, [phi]II increased by as much as 115% in P. bisulcatum but by no more than 17% in P. antidotale. Under the same conditions [phi]II increased to an intermediate degree in P. milioides. Measurements of accumulation of the photooxidized form of the photosystem I reaction center (P700+) based on the light-dependent in vivo absorbance change at 830 nm indicate that the steady-state concentration of P700+ varied in an antiparallel manner with [phi]II when either the irradiance or O2 concentration was changed. Hence, O2-dependent changes in [phi]II were indicative of variations in linear photosynthetic electron transport. These experiments revealed, however, that a significant capacity was retained for in vivo regulation of the apparent quantum yield of photosystem I ([phi]I) independently of [phi]II+ Coordinate regulation of quantum yields of photosystems I and II (expressed as [phi]I:[phi]II in response to changing irradiance and O2 level differed markedly for the C3 and C4 species, and the response for the C3-C4 species most closely resembled that observed for the C4 species. The fraction of total linear electron transport supporting photorespiration at 213 mbars of O2 was negligible in the C4 species and was 13% lower in the C3-C4 species relative to the C3 species as calculated from fluorescence and gas-exchange determinations. At high photon-flux rates and high O2 concentration, the potential benefit to light use for net CO2 uptake arising from lower photorespiration in P. milioides was offset by a reduced capacity for total CO2- and O2-dependent noncyclic electron transport in this species compared with P. bisulcatum.     

Co-function of C3-and C4-photosynthetic pathways in C3, C4 and C3-C4 intermediate Flaveria species

The potential for C₄ photosynthesis was investigated in five C₃-C₄ intermediate species, one C₃ species, and one C₄ species in the genus Flaveria, using ¹⁴CO₂ pulse-¹²CO₂ chase techniques and quantum-yield measurements. All five intermediate species were capable of incorporating ¹⁴CO₂ into the C₄ acids malate and aspartate, following an 8-s pulse. The proportion of ¹⁴C label in these C₄ products ranged from 50–55% to 20–26% in the C₃-C₄ intermediates F. floridana Johnston and F. linearis Lag. respectively. All of the intermediate species incorporated as much, or more, ¹⁴CO₂ into aspartate as into malate. Generally, about 5–15% of the initial label in these species appeared as other organic acids. There was variation in the capacity for C₄ photosynthesis among the intermediate species based on the apparent rate of conversion of ¹⁴C label from the C₄ cycle to the C₃ cycle. In intermediate species such as F. pubescens Rydb., F. ramosissima Klatt., and F. floridana we observed a substantial decrease in label of C₄-cycle products and an increase in percentage label in C₃-cycle products during chase periods with ¹²CO₂, although the rate of change was slower than in the C₄ species, F. palmeri. In these C₃-C₄ intermediates both sucrose and fumarate were predominant products after a 20-min chase period. In the C₃-C₄ intermediates, F. anomala Robinson and f. linearis we observed no significant decrease in the label of C₄-cycle products during a 3-min chase period and a slow turnover during a 20-min chase, indicating a lower level of functional integration between the C₄ and C₃ cycles in these species, relative to the other intermediates. Although F. cronquistii Powell was previously identified as a C₃ species, 7–18% of the initial label was in malate+aspartate. However, only 40–50% of this label was in the C-4 position, indicating C₄-acid formation as secondary products of photosynthesis in F. cronquistii. In 21% O₂, the absorbed quantum yields for CO₂ uptake (in mol CO₂·[mol quanta]^-1) averaged 0.053 in F. cronquistii (C₃), 0.051 in F. trinervia (Spreng.) Mohr (C₄), 0.052 in F. ramosissima (C₃-C₄), 0.051 in F. anomala (C₃-C₄), 0.050 in F. linearis (C₃-C₄), 0.046 in F. floridana (C₃-C₄), and 0.044 in F. pubescens (C₃-C₄). In 2% O₂ an enhancement of the quantum yield was observed in all of the C₃-C₄ intermediate species, ranging from 21% in F. ramosissima to 43% in F. pubescens. In all intermediates the quantum yields in 2% O₂ were intermediate in value to the C₃ and C₄ species, indicating a co-function of the C₃ and C₄ cycles in CO₂ assimilation. The low quantum-yield values for F. pubescens and F. floridana in 21% O₂ presumably reflect an ineffcient transfer of carbon from the C₄ to the C₃ cycle. The response of the quantum yield to four increasing O₂ concentrations (2–35%) showed lower levels of O₂ inhibition in the C₃-C₄ intermediate F. ramosissima, relative to the C₃ species. This indicates that the co-function of the C₃ and C₄ cycles in this intermediate species leads to an increased CO₂ concentration at the site of ribulose-1,5-bisphosphate carboxylase/oxygenase and a concomitant decrease in the competitive inhibition by O₂.Abbreviations PEP phosphoenolpyruvate - PGA 3-phosphoglycerate - RuBP ribulose-1,5-bisphosphate     

Photosynthesis, Morphology, Leaf Anatomy, and Cytogenetics of Hybrids between C(3) and C(3)/C(4)Panicum Species

The Laxa group of the Panicum genus contains species which have CO₂ exchange and anatomical characteristics intermediate to C₃ and C₄ photosynthetic types (C₃/C₄), and also species characterized as C₃. Hybrids were made between two of the C₃/C₄ species and two C₃ species. Carbon dioxide exchange and morphological, leaf anatomical, and cytogenetic characteristics of F₁ hybrids between Panicum milioides Nees. ex Trin (C₃/C₄) and P. laxum Mez. (C₃), P. spathellosum Doell (C₃/C₄) and P. boliviense Hack. (C₃), and P. spathellosum and P. laxum were studied. There were no consistent differences in apparent photosynthesis, although two of the three hybrids had higher net CO₂ uptake than the C₃ parent. Values of inhibition of apparent photosynthesis by 21% O₂, CO₂ loss in the light, and CO₂ compensation concentration for the hybrids were between those of the parents. All three hybrids showed leaf anatomical traits, especially organelle quantities in the bundle sheath cells, between those of their respective parents. Linear regression of CO₂ compensation concentration on the percentage of mitochondria and chloroplasts in vascular bundle sheaths of the parents and hybrids gave correlation coefficients of −0.94. This suggests that the reduction in CO₂ loss in the C₃/C₄ species, and to a lesser degree in the F₁ hybrids, was due to development of organelles and perhaps a higher proportion of leaf photorespiration in bundle sheaths. The overall morphology of the hybrids was so different from the parents that they could be described as new taxonomic forms. The chromosomes in the hybrids were mainly unpaired or paired as bivalents indicating possible homology between some parental genomes.     

Germination Shifts of C3 and C4 Species under Simulated Global Warming Scenario

Research efforts around the world have been increasingly devoted to investigating changes in C₃ and C₄ species'' abundance or distribution with global warming, as they provide important insight into carbon fluxes and linked biogeochemical cycles. However, changes in the early life stage (e.g. germination) of C₃ and C₄ species in response to global warming, particularly with respect to asymmetric warming, have received less attention. We investigated germination percentage and rate of C₃ and C₄ species under asymmetric (+3/+6°C at day/night) and symmetric warming (+5/+5°C at day/night), simulated by alternating temperatures. A thermal time model was used to calculate germination base temperature and thermal time constant. Two additional alternating temperature regimes were used to test temperature metrics effect. The germination percentage and rate increased continuously for C₄ species, but increased and then decreased with temperature for C₃ species under both symmetric and asymmetric warming. Compared to asymmetric warming, symmetric warming significantly overestimated the speed of germination percentage change with temperature for C₄ species. Among the temperature metrics (minimum, maximum, diurnal temperature range and average temperature), maximum temperature was most correlated with germination of C₄ species. Our results indicate that global warming may favour germination of C₄ species, at least for the C₄ species studied in this work. The divergent effects of asymmetric and symmetric warming on plant germination also deserve more attention in future studies.     

Epidermal Patterning and Silica Phytoliths in Grasses: An Evolutionary History

Due to the immense ecological and economic significance of grasses, their highly characteristic long–short epidermal patterning and associated silica phytoliths represent significant diagnostic markers in studies of ancient climate change and agriculture. We explore the link between epidermal cell patterning and phytolith development and review the evolutionary history of phytoliths in the context of recent well-resolved phylogenetic analyses of grasses and allied Poales, focusing on early-divergent grasses and the subfamilies that constitute the BEP group (the bamboos and their allies). Dimorphic epidermal patterning is a common feature of Poaceae and the related family Joinvilleaceae, where phytoliths are located primarily in the short cells. However, Joinvillea lacks the short-cell pairs that occur in many grasses. The costal rows of phytoliths that characterize some grasses could represent loss of long–short cell patterning over the veins. Unlobed phytoliths probably represent the ancestral condition in grasses, though bilobate phytoliths evolved at an early stage. Either transverse-unlobed or transverse-bilobate phytoliths predominate in the early-divergent lineages, whereas axial-bilobates (or polylobates) primarily characterize the PACMAD clade and the BEP subfamily Pooideae.     

C3、C4和C3-C4中间型植物的进化

介绍了有关C3、C4和C3-C4中间型植物进化的形态学、生理学、分子生物学、遗传学等方面的证据;推断地球上首先出现C3植物,然后是C3-C4中间类型植物,最后出现C4植物.     

Photosynthesis, Leaf Anatomy, and Morphology of Progeny from Hybrids between C(3) and C(3)/C(4)Panicum Species

Species in the Laxa group of Panicum have C₃ or C₃/C₄ photosynthesis based on leaf anatomical and CO₂ exchange characteristics. Hybrids were previously made between C₃/C₄ and C₃ species in this group (RH Brown et al. 1985 Plant Physiol 77: 653-658). In this paper, CO₂ exchange, morphological, and leaf anatomical characteristics of F₂ or F₅ progeny from colchicine-induced amphiploids of C₃/C₄ × C₃ hybrids (Panicum milioides Nees ex Trin. [C₃/C₄] × Panicum laxum Mez [C₃] and Panicum spathellosum Doell [C₃/C₄] × Panicum boliviense Hack. [C₃]) were studied.

There were no differences found in morphology or physiology between the amphiploids and the F₁ hybrids from which they were produced. In the segregating progeny, CO₂ compensation concentration and photorespiration values typical of C₃, but not of C₃/C₄ plants, were recovered. Progeny were found from both crosses which possessed O₂ inhibition of apparent photosynthesis typical of the parents, and in the case of the P. milioides × P. laxum cross, leaf anatomy and overall plant morphology typical of the parents were observed in some progeny. The progeny were found to possess recombinations of various traits associated with reduced photorespiration, so that no correlation existed among O₂ inhibition of apparent photosynthesis, CO₂ compensation concentration, and leaf anatomical traits. One plant was especially noteworthy in possessing leaf anatomy typical of C₃/C₄ plants, but with CO₂ exchange characteristics of C₃ plants.