The effects of clipping height and frequency on net primary production of Andropogon gerardii (C4 grass) and Bromus inermis (C3 grass) in greenhouse experiments

There are potential agronomic and environmental benefits from incorporating warm‐season (C₄) grasses into temperate pasture systems, usually dominated by cool‐season (C₃) grasses, but there is a lack of information on how frequency and height of defoliation affects C₄ grasses. Three greenhouse experiments were conducted under (i) spring, (ii) summer and (iii) spring + summer clipping regimes. In each experiment, the effects of clipping frequency (weekly and monthly) and clipping height (clipped to 5 and 10 cm) were determined on above‐ and below‐ground net primary production (ANPP and BNPP) and total and seasonal dry matter (DM) yield for Andropogon gerardii Vitman (big bluestem, C₄ grass) and Bromus inermis Leyss (smooth brome, C₃ grass). Six replicates per treatment were used. In all experiments, ANPP and BNPP of smooth brome was greater than that of big bluestem although during late summer months big bluestem had higher DM yields of herbage than smooth brome. There were different effects of frequency and height of clipping for both species on two similar measurements: total annual DM yield and ANPP, indicating that the ability to generalize about the effects of defoliation from ecological and agronomic grassland standpoints is questionable. Clipping effects on ANPP and BNPP were different for summer‐clipped pots than for spring, and spring + summer‐clipped pots, indicating that management could be tailored to meet specific agronomic or conservation goals.     

Seed production in grassland species: Morpho‐biological determinants in a species‐rich semi‐natural grassland

Studying the seed production of herbaceous species can help to conserve grassland habitats and re‐create new high‐value grassland surfaces. Studies on grassland seed production have focused mainly on individual species and traits, without characterizing their relative importance at the plant community level. The aim of this study was to investigate the entire seed production process of the main species in a temperate grassland. Fertile shoots (FS) of twenty‐nine grasses and forbs were collected over 4 years and analysed for sixteen traits that determine inflorescence size, seed production and seed quality. The per cent viability played a predominant role in determining the total production of viable seeds. Forbs showed a range of reproductive strategies, including variable distribution of flowers among growth periods, number of inflorescences per FS and relationship between seed size and FS density in the grassland. The flower production for grasses was concentrated in the first growth period, but this limitation was mitigated by a higher seed dormancy. The number of viable seeds per FS and seed size were important components of the reproductive strategy of forbs, with heavy‐seeded species being characterised by high individual densities in the community, but producing few seeds per FS. Light‐seeded species showed an opposite pattern. The results suggest that when using seeds from semi‐natural grasslands for ecological restoration, special attention should be paid to the seed amount, germinability and viability of forbs, as they seem to depend more on seed reproduction and have a lower ovule to seed transformation efficiency.     

Variability in grass forage quality and quantity in response to elevated CO2 and water limitation

Global climate change is expected to alter carbon dioxide concentration ([CO₂]) and water availability, with uncertain impacts on agriculture. Forage quality and quantity in grazing systems are of particular concern because C₃ and C₄ plants respond differently to altered environmental conditions. In a growth chamber, we compared crude protein content, biomass recovery and total crude protein across a set of perennial C₃ and C₄ grasses from the northern U.S. Great Plains under elevated [CO₂] and simulated drought. Simulated 95% confidence intervals indicate both C₃ and C₄ grasses increased forage quality and quantity under elevated [CO₂]. C₄ grasses were generally resistant to water limitation while forage quality and quantity of C₃ grasses declined under simulated drought. Our results are consistent with literature on forage quantity responses to elevated [CO₂] and drought, but forage quality responses contradict expectations. We suggest measuring plant functional traits might better elucidate response mechanisms and ameliorate methodological differences even if traits are not directly applicable to grazing management.     

Plant diversity effects on herbage production and compositional changes in New Zealand hill country pastures

A small‐plot field experiment on grazed hill country pastures in the North Island of New Zealand was conducted to examine the productivity and compositional characteristics of swards in response to variation in pasture species diversity. The balanced incomplete factorial design incorporated variation in location, slope, soil fertility and combinations of eight plant functional groups (C₄ grasses, annual grasses, annual legumes, perennial C₃ grasses, perennial legumes, perennial forbs, ryegrass and browntop). Net herbage accumulation and botanical composition were measured at 18 months (spring) and 24 months (autumn) after oversowing following application of a systemic herbicide. Analysis of variance indicated a significant positive relationship between the number of functional groups sown and herbage accumulation of the sown species in spring, but not with total herbage accumulation. Regression analysis showed that herbage accumulation was also affected by the identity of the functional groups. However, the statistical models indicated that pasture productivity was most strongly influenced by site factors. There was a significant negative relationship between both the number and herbage accumulation of unsown species and the number of functional groups sown, indicating a positive relationship between diversity and resistance to invasion by unsown species. A comparison of the vegetation between the plots before and after oversowing showed that those more diverse prior to sowing returned to their initial composition more rapidly, evidence that diverse vegetation was more resilient in the face of disturbance.     

Potential of the indigenous desert grasses of the Arabian Peninsula for forage production in a water‐scarce region

Indigenous perennial grasses are widely distributed in the Arabian Peninsula. Their survival under limited rainfall and grazing suggests a potential role as grassland species and for rehabilitation of degraded rangelands. Forage productivity, seed production and water‐use efficiency (WUE) was determined over 2 years for four indigenous grasses: buffel grass (Cenchrus ciliaris L.), dakhna (Coelachyrum piercei Benth.), da’ay (Lasiurus scindicus Henr.) and tuman (Panicum turgidum Forssk.) together with one exotic species, rhodes grass (Chloris gayana Kunth) in the central region of the United Arab Emirates. Three irrigation treatments were used: R1 (1858–6758 m³ ha^?1 year^?1), R2 (929–3379 m³ ha^?1 year^?1) and R3 (464–1689 m³ ha^?1 year^?1). Buffel grass had the highest dry‐matter (DM) yield under all irrigation treatments. The average DM yield of buffel grass was 14·6 and 15·1 t ha^?1 in the 2 years which was significantly higher than that for the other grasses with dakhna having the lowest DM yields. The WUE of 0·7 and 0·8 kg DM m^?3 in the 2 years for buffel grass was significantly greater than for the other grasses. Buffel grass showed the highest increase in WUE in both years when the irrigation was reduced from treatment R1 to R3. The results suggest that the desert grasses of the Arabian Peninsula, such as buffel grass, could be useful grass species in reducing the use of scarce irrigation water provided that seed production can be increased.     

Variations in structural,biochemical, and physiological traits of photosynthesis and resource use efficiency in Amaranthus species (NAD-ME-type C4)

C₄ plants show higher photosynthetic capacity and productivity than C₃ plants owing to a CO₂-concentrating mechanism in leaves, which reduces photorespiration. However, which traits regulate the photosynthetic capacity of C₄ plants remains unclear. We investigated structural, biochemical, and physiological traits associated with photosynthesis and resource use efficiency in 20 accessions of 12 species of Amaranthus, NAD-malic enzyme-type C₄ dicots. Net photosynthetic rate (P_N) ranged from 19.7 to 40.5 μmol m^?2 s^?1. P_N was positively correlated with stomatal conductance and nitrogen and chlorophyll contents of leaves and was weakly positively correlated with specific leaf weight. P_N was also positively correlated with the activity of the C₃ enzyme ribulose-1,5-bisphoshate carboxylase/oxygenase, but not with the activities of the C₄ enzymes phosphoenolpyruvate carboxylase and NAD-malic enzyme. Structural traits of leaves (stomatal density, guard cell length, leaf thickness, interveinal distance, sizes of mesophyll and bundle sheath cells and the area ratio between these cells) were not significantly correlated with P_N. These data suggest that some of the biochemical and physiological traits are involved in interspecific P_N variation, whereas structural traits are not directly involved. Photosynthetic nitrogen use efficiency ranged between 260 and 458 μmol mol^?1 N s^?1. Photosynthetic water use efficiency ranged between 5.6 and 10.4 mmol mol^?1. When these data were compared with previously published data of C₄ grasses, it is suggested that common mechanisms may determine the variations in resource use efficiency in grasses and this dicot group.     

Seasonal productivity and nutritive value of temperate grasses found in semi-natural pastures in Europe: responses to cutting frequency and N supply

Monocultures of thirteen perennial C₃ grass species that co‐occur in temperate semi‐natural grassland communities in Europe were compared in a factorial field design of two levels of N supply and two levels of cutting frequency. Above‐ground yield of dry matter (DM), crude protein (CP) concentration and pepsin‐cellulase DM digestibility of herbage were measured in two successive years. Species was the largest source of variability in yield of DM and DM digestibility of herbage, while CP concentration of herbage responded more to management factors. The highest mean DM yields and values of DM digestibility of herbage were achieved in spring for Festuca arundinacea and in autumn for Phleum pratense. Poa trivialis and Festuca rubra had the lowest DM yield and DM digestibility values, respectively, regardless of seasons and treatments. For all species CP concentration in herbage increased in response to an increase in cutting frequency and N supply by an average of 46 and 34 g kg^?1 DM respectively. Differences between years and seasons indicated the importance of plant phenology on nutritional variables and the influence of environmental factors on species performance. Species ranking was compared according to their annual digestible DM and CP yields. The results show that some grasses have a nutritive value which is comparable to that of forages selected for high yields.     

广东雷州杂草稻sh4和qSH1基因的功能SNP位点序列分析

sh4和qSH1基因是水稻的主效落粒基因,本研究旨在测定雷州杂草稻的sh4和qSH1的基因型,为雷州杂草稻的落粒机制研究提供分子基础。本研究从雷州10个杂草稻群体采集了100个单株及10份栽培稻种子,观测其颖壳颜色、果皮颜色和芒及落粒率,并对sh4和qSH1的功能SNP位点进行PCR扩增及序列分析。结果表明,57份杂草稻的sh4功能SNP位点为野生落粒型（G）,15份为杂合型G/T,28份为突变难落粒型（T）。其中,G型和G/T型的72份杂草稻的落粒率都极高（95%~100%）,而28份T型杂草稻中,19份为中度落粒（30%~75%）,5份杂草稻的落粒率很高（>95%）,4份杂草稻的落粒率小于30%。100份杂草稻和10份栽培稻的qSH1功能SNP位点均为G。结合落粒表型及基因型,推测sh4是调控雷州杂草稻落粒性的一个主要基因。     

Comparative defoliation tolerance of temperate perennial grasses

The defoliation tolerance of cultivars of four temperate perennial pasture grasses, perennial ryegrass (Lolium perenne, cv. Yatsyn1), phalaris (Phalaris aquatica cv. Australian), tall fescue (Festuca arundinaceae cv. Demeter) and cocksfoot (Dactylis glomerata cv. Porto), was determined under controlled conditions over a period of 12 weeks. Undefoliated plants were compared with defoliated plants, where only half of one leaf was left intact at the initial defoliation, and leaf regrowth was harvested every 3–4 d. The growth responses measured were plant tiller number, dry weight, relative leaf regrowth rate, root:shoot ratio, sheath:stem ratio and specific leaf weight. All species showed morphological adaptations that potentially increased their ability to tolerate defoliation (e.g. increased allocation to shoot at the expense of roots and lower specific leaf weights) but cocksfoot was found to be the most defoliation‐tolerant and perennial ryegrass the least. The adaptation that favoured cocksfoot most strongly was high sheath:stem ratio, which, it is proposed, allowed it to maintain photosynthesis and a level of carbon supply sufficient to support regrowth throughout the experiment. The strategy of perennial ryegrass which favours leaf growth and leads to rapid leaf turnover rates made it particularly susceptible to defoliation under the conditions of this experiment. This highlights the likely importance of defoliation‐avoidance responses in explaining the well‐known grazing resistance of this species. Phalaris and tall fescue showed responses that were intermediate between the other two species. The importance of defoliation‐avoidance mechanisms and implications for grazing management are discussed.     

Impacts of dormancy‐regulating chemicals on innate and salinity‐induced dormancy of four forage grasses native to Arabian deserts

The effects of dormancy‐regulating chemicals (DRCs) on alleviating innate and salinity‐induced dormancy (SID) were assessed in seeds of four perennial forage grasses (Pennisetum divisum, Sporobolus spicatus, Coelachyrum brevifolium and Centropodia forsskalii). These grasses have the potential to be used for restoration or rehabilitation of degraded rangelands of the Arabian Gulf deserts. The four species showed various levels of innate dormancy. Germination of seeds stored for 2 months was not enhanced by any of the five studied DRCs in both C. brevifolium and P. divisum, but significantly improved by thiourea, fusicoccin and gibberellic acid (GA) in C. forsskalii and by thiourea, fusicoccin and nitrate in S. spicatus. Salinity had a significant negative effect on all the studied grasses. Sporobolus spicatus was more tolerant to salinity, compared with the other species. The effect of DRCs on alleviating SID depended on species. Whereas DRCs had little effect on alleviating SID in C. forsskalii and P. divisum, they greatly alleviated it in S. spicatus and C. brevifolium. Partial alleviation was observed by fusicoccin in C. brevifolium and by GA, kinetin and thiourea in C. forsskalii. Nitrate, fusicoccin and GA resulted in a complete alleviation in S. spicatus seeds in 200 mm NaCl.     

Ultrastructure of Fusion Product between Protoplasts from C3 and C4 Species of Amaranthaceae

Summary

This paper describes the ultrastructure of the electric field-induced fusion products of C₃ and C₄ species of Amaranthaceae at the early developmental stage. Protoplasts of C₃ species were isolated from a Ceiosia cristata L. cell suspension and, those of C₄ species were isolated from an Amaranthus tricolor L. cotyledon. Incompatibility occurred in the C₃/C₄ hybrid. The incompatibility reactions were detected in the newly formed hybrid cells accompanied with significant changes in the nucleolus (segregation of nucleolar components) and plastids (cup-like shape or amoeboid plastid enclosing cytoplasmic materials) of C₃ species parent. The structural changes in the organelles of the C₄ partner were less marked. After 5 days of culture, most organelles showed high cellular activity, and a normal dedifferentiation process of mesophyll chloroplasts was observed. At this stage nucleolar segregation was not detected and the C₃ species plastids were difficult to distinguish from the proplastids formed from mesophyll chloroplasts. In addition, some mitochondria showed bursting-like structure. However, under the culture condition used these somatic incompatibility did not seem to impair further growth of fusion products since they were still proliferating well resulting in callus formation.     

Seed dormancy and dormancy‐breaking methods in Leymus chinensis (Trin.) Tzvel. (Poaceae)

Leymus chinensis (Trin.) Tzvel. is a perennial grass with high productivity and forage value; however, poor stand establishment, often due to seed dormancy, limits its widespread use for forage production. To investigate the mechanism of seed dormancy and to develop effective methods of improving germination, the contribution of each part of the caryopsis to dormancy was investigated, and a number of single or combined dormancy‐breaking pre‐treatments were conducted using three seed lots. The palea, lemma, pericarp/testa, and endosperm all contributed to seed dormancy. The contribution of each part to dormancy was 23·4%, lemma; 6·2%, palea; 28·4%, pericarp/testa; and 42·0%, endosperm. Hull (palea and lemma) removal, pericarp/testa piercing, and soaking in distilled water or 30% sodium hydroxide (NaOH) significantly decreased the percentage of dormant seeds (i.e. increased germination). Treating hull‐removed and pericarp/testa‐pierced seeds with gibberellic acid (GA₃) also significantly decreased the percentage of dormant seeds. Compared with each of the single pre‐treatments, the combined pre‐treatment of pre‐soaking in water for 1 d, then 30% NaOH for 60 min and treating with 300 μm GA₃ resulted in the highest germination (89%); and seed viability was 91%.     

Impact of deferred grazing and fertilizer on herbage production,soil seed reserve and nutritive value of native pastures in steep hill country of southern Australia

Developing sustainable grazing management systems based on perennial species is critical to preventing land degradation in marginal land classes. A field study was conducted from 2002 to 2006 to identify the impacts of deferred grazing (no defoliation of pastures for a period generally from spring to autumn) and fertilizer application on herbage accumulation, soil seed reserve and nutritive value in a hill pasture in western Victoria, Australia. Three deferred grazing strategies were used: short‐term deferred grazing (no defoliation between October and January), long‐term deferred grazing (no defoliation from October to the autumn break) and optimized deferred grazing (withholding time from grazing commenced between annual grass stem elongation and seed head emergence and concluded in February/March). These treatments were applied with two fertilizer levels (with or without fertilizer at 50 kg phosphorus ha^?1 and 2000 kg lime ha^?1 applied in year 1 only) in a factorial arrangement and two additional treatments: continuous grazing (CG) and no grazing (NG) in year 1. The deferred grazing treatments on average produced herbage dry matter of 4773 kg ha^?1, the NG produced 4583 kg ha^?1 and the CG produced 3183 kg ha^?1 in year 4 (2005–06) of the experiment. Deferred grazing treatments with and without fertilizer application produced an average of 5135 and 4411 kg DM ha^?1 respectively. Averaged over 4 years, deferred grazing increased the germinable seed pool of perennial grasses by 200% and annual grasses by 50% (except optimized deferred grazing that considerably decreased the annual grass seed pool) compared with the CG. The best of the deferred grazing strategies increased the digestibility of pastures by 7% compared with the CG. The results demonstrated that deferred grazing from spring to autumn followed by rotational grazing could be an effective tool to increase herbage production and soil seed pool and improve the digestibility of native pastures in the steep hill country of southern Australia.     

THE INDUCTIVE REQUIREMENTS FOR FLOWERING OF SOME TEMPERATE GRASSES

A survey was made of the flowering responses of 47 annual, biennial and perennial grasses, to measure the extent of any winter requirement for floral induction, and to see how far this could be provided by controlled cold or short-day treatment.
The annual species showed little or no inductive requirement, nor did the perennials Arrheuatherum elatius, Ceratochloa unioloides, Phleum pratense and Poa nemoralis . Most perennials possessed a definite inductive requircinent. A few species, such as Lolium perenne , responded to both cold and short-day given to the young seedling, while others, such as Phleum nodosum, Agrostis alba, A. canina, A. stolonifera and A. tenuis , responded to short-day induction but not to cold.
Many temperate perennials, however, including Cynosurus cn'status, Dactylis glomerata, Festuca pratensis, F. rubra, Poa pratensis and P. trivialis did not respond to seedling induction and possibly have a juvenile stage before they are able to respond to inductive conditions.
The sequence of flowering responses in the temperate perennial grasses is evidently more complex than was previously thought, and the perennial habit can be achieved by many different developmental pathways.     

Water use efficiency and drought survival in Mediterranean perennial forage grasses

In Mediterranean areas, water use efficiency (WUE) is mainly increased by maximising crop growth during the rainy seasons. Perennial forage species have a number of advantages in comparison to the predominantly used annuals. They can utilize water from autumn to spring, while annuals need to be sown or to germinate from the soil seed bank. Under Mediterranean annual rainfall pattern, perennial plants must grow from autumn to spring and survive under summer aridity. Drought survival can impact water use efficiency through plant mortality and stand recovery after autumn rainfalls. In order to enhance knowledge of physiological and agronomic traits associated with WUE and persistence, a 3-year study was conducted at two Mediterranean sites, comparing a range of accessions of two perennial species, cocksfoot (Dactylis glomerata L.) and tall fescue (Lolium arundinaceum (Schreb.) Darbysh). Within the accessions of predominantly Mediterranean origin, we defined four major functional types, i.e. groups of accessions with similar response to summer drought. One type (FT1) is represented by the only cultivar of a semi-arid cocksfoot (Kasbah) that is completely summer dormant, with high persistence under the most arid situations but with low WUE. The type FT2 all cocksfoot cultivars (Currie, Delta-1, Jana, Medly, and Ottava) that are not or less summer dormant, with good perenniality and intermediate productivity at rainy seasons. The type FT3 includes the cultivars of tall fescue (Centurion, Flecha, Fraydo, and Tanit) that combine an incomplete summer dormancy, a deep rooting system and the highest WUE from autumn to spring. The type FT4 is represented by a cultivar of tall fescue (Sisa) with no summer dormancy, therefore less persistent and also less productive. Dehydration avoidance in tall fescue and cocksfoot and summer dormancy in cocksfoot were the main strategies contributing to persistence under summer drought. WUE in autumn was highly correlated with sward recovery after drought. Seasonal and total WUE were also highly correlated with biomass production over the same period and with depth and density of the root system. Parameterization of functional types of the major species of forage grasses will enhance future modelling work to test the effects of a range of environments and future climate scenarios.     

Effects of nitrogen fertilization and cutting intensity on the agronomic performance of warm‐season grasses

Responses of grasses to N fertilization are affected by cutting intensity although little is known regarding the interactions of these factors in warm‐season grasses. Pre‐cutting canopy height, herbage accumulation and changes in the nutritive value of warm‐season grasses in response to four different management strategies were assessed from October 2011 to September 2014. Treatments included two cutting intensities (70 vs. 50% depletion of canopy height set by 95% light interception), two N fertilization levels (zero vs. 300 kg N ha^?1 year^?1) and six perennial C₄ grass species (Axonopus catharinensis; Cynodon spp. hybrid Tifton 85; Hemarthria altissima cv. Flórida; Megathyrsus maximus cv. Aruana; Paspalum notatum cv. Pensacola; and Urochloa brizantha cv. Marandu) grown in monoculture in a factorial experimental design. Canopy height varied among grass species, cutting intensity and N treatments, mainly among seasons, indicating that more than one management target (i.e. canopy height) existed throughout the plant growth cycle for each species. The largest herbage accumulation occurred in the N fertilization treatments for most species, regardless of cutting intensity. Nitrogen fertilization and 50% depletion of canopy height increased the leaf proportion and decreased the neutral detergent fibre content. Overall, N fertilization had a stronger positive impact than cutting intensity on the acid detergent fibre content, dry‐matter digestibility and crude protein content, but the magnitudes of the responses were species‐specific.     

Breeding perennial grasses for forage usage: An experimental assessment of trait changes in diploid perennial ryegrass (Lolium perenne L.) cultivars released in the last four decades

Breeding to improve perennial grasses for forage usage aims to increase the whole-plant aerial biomass. However, the ability to increase the whole-plant yield by breeding may be questionable in grasses which are expected to reach an optimal leaf area index during grass stand growth rather rapidly. We carried out a field experiment aiming to compare herbage yield and other performances of seven natural populations and 21 cultivars of diploid perennial ryegrass registered on European National lists in the last 40 years. Cultivars were sampled in the spike emergence earliness range used for pasture usage, i.e. in the semi-late and late earliness range according to the French classification into spike emergence earliness groups. Forage performances of populations and cultivars were tested in four locations in monthly cut dense sward plots. Morphological and phenological traits were recorded in two spaced-plant experiments, and seed yield was assessed in two locations in standard seed production conditions. Trait regression on cultivar registration year pointed out that total dry-matter yield of cultivars increased by +3.2% per decade. However, dry-matter yield was not improved in the most favourable period for grass growth, i.e. in spring. On the other hand, it was improved in summer (+2.8% per decade) and autumn (+7.4% per decade) when the aerial biomass production is on average rather small. Breeders succeeded in significantly reducing aftermath heading. Reduction in aftermath heading is likely to have contributed to the noted increase in summer and autumn dry-matter yields, and to a moderate improvement in herbage feeding value. Breeding was also efficient in improving rust resistance and in bringing forward the start of spring growth. For monthly cut swards, we evidenced a clear association of leaf and lamina lengths with spring and summer dry-matter yields. Breeding for long leaves, or for high leaf elongation rate, should contribute to improve the cumulated intercepted radiation during re-growths, and consequently could be a way of improving the spring dry-matter yield. Seed yield did not improve with breeding for forage performances. However, we did not notice any negative association between the seed yield and traits related with forage usage. Direct selection pressures on seed yield criteria applied at the core of the breeding process should enable to improve the seed yield without any negative impact on forage performances.     

Intracellular position of mitochondria and chloroplasts in bundle sheath and mesophyll cells of C3 grasses in relation to photorespiratory CO2 loss

In C₃ plants, photosynthetic efficiency is reduced by photorespiration. A part of CO₂ fixed during photosynthesis in chloroplasts is lost from mitochondria during photorespiration by decarboxylation of glycine by glycine decarboxylase (GDC). Thus, the intracellular position of mitochondria in photosynthetic cells is critical to the rate of photorespiratory CO₂ loss. We investigated the intracellular position of mitochondria in parenchyma sheath (PS) and mesophyll cells of 10 C₃ grasses from 3 subfamilies (Ehrhartoideae, Panicoideae, and Pooideae) by immunostaining for GDC and light and electron microscopic observation. Immunostaining suggested that many mitochondria were located in the inner half of PS cells and on the vacuole side of chloroplasts in mesophyll cells. Organelle quantification showed that 62–75% of PS mitochondria were located in the inner half of cells, and 62–78% of PS chloroplasts were in the outer half. In mesophyll cells, 61–92% of mitochondria were positioned on the vacuole side of chloroplasts and stromules. In PS cells, such location would reduce the loss of photorespiratory CO₂ by lengthening the path of CO₂ diffusion and allow more efficient fixation of CO₂ from intercellular spaces. In mesophyll cells, it would facilitate scavenging by chloroplasts of photorespiratory CO₂ released from mitochondria. Our data suggest that the PS cells of C₃ grasses have already acquired an initial structure leading to proto-Kranz and further C₃–C₄ intermediate anatomy. We also found that in the Pooideae, organelle positioning in PS cells on the phloem side resembles that in mesophyll cells.     

Photosynthetic Characteristics of an Amphibious C4 Plant,Eleocharis retroflexa ssp. chaetaria

Abstract

Eleocharis retroflexa (Poir.) Urban ssp. chaetaria (Roem. & Schult.) T. Koyama, an amphibious leafless sedge, grows not only under terrestrial conditions but also under completely submerged aquatic conditions. We investigated the photosynthetic traits and structural features of the culms, which are the photosynthetic organs, in the terrestrial and submerged forms of this species and compared them with those of other amphibious species of Eleocharis which are known to change the photosynthetic modes. The culms of the terrestrial form had Kranz anatomy with well-developed Kranz (bundle sheath) cells and high levels of C₄ enzyme activity typical of the NAD-malic enzyme (NAD-ME) subtype of C₄ metabolism. They also had a δ ¹³C value typical of C₄ plants, indicating that the terrestrial form fixes carbon through the C₄ pathway. The culms of the submerged form had not only a Kranz-like anatomy but also revealed anatomical traits typical of leaves of submerged aquatic plants. The activities of the C₄ enzymes in the submerged form were lower than those in the terrestrial form, but were still in the range typical of G4 plants, ¹⁴C pulse-¹²C chase experiments with the submerged form indicated that almost all of the fixed ¹⁴C was incorporated into G4 compounds, and subsequently the raioactivity was transferred into C₃ compounds and sucrose. The submerged form showed no diurnal fluctuation in malate level. These data demonstrate that a C₄ metabolism is operative even in the submerged form. This unique amphibious C₄ plant provides an intriguing example of the physiological and ecological adaptability of C₄plants.     

Life strategies and spatial arrangement of grasses in a Mediterranean ecosystem in Greece

The temporal and spatial distribution, and the life strategies, of fifteen grass species in a Mediterranean evergreen sclerophyllous formation at Petralona (Chalkidiki, Greece) were studied in order to identify the relationships between species, and among species and the environment, which allows for the coexistence of species. The grass species were grouped into (i) early annual species that were characterized by early germination and a relatively long growth period; they did not need excessive resources and were distributed over the entire site, (ii) late annuals that were characterized by late germination and a short growth period; they required resources within this limited period and were competitive in capturing them and so were confined to the most productive sites of the site and (iii) perennial grasses that were characterized by early germination and vegetative reproduction; they were tolerant to nutrient shortages and hence maintained their distribution on the site. The spatial arrangement of species can be considered as a response to the selective pressures of the seasonal water and resource availability in a Mediterranean environment. This environment posed temporal limitations on the activity of early annual and late annual species, which restricts them in time and space. The perennial species overcame this by being flexible in their resource capture and use, because of their ability to store nutrients.