A dynamic analysis of the shade-induced plasticity in Arabidopsis thaliana rosette leaf development reveals new components of the shade-adaptative response

BACKGROUND AND AIMS: It is well known that plant aerial development is affected by light intensity in terms of the date of flowering, the length of stems and petioles, and the final individual leaf area. The aim of the work presented here was to analyse how shade-induced changes in leaf development occur on a dynamic basis from the whole rosette level to that of the cells. METHODS: Care was taken to ensure that light intensity was the only source of micro-meteorological variation in the study. The dynamics of leaf production, rosette expansion, individual leaf area expansion and epidermal cell expansion were analysed in Arabidopsis thaliana plants grown under two light intensities in three independent experiments. KEY RESULTS: The total area of rosette leaves was reduced by the shading treatment. Both the number of leaves produced and their individual leaf areas were reduced. The reduction in leaf number was associated with a reduction in leaf initiation rate and the duration of the phase of leaf production. The reduction in individual leaf area was associated with a reduction in leaf expansion rate and an increase in the duration of leaf expansion. The changes in leaf expansion dynamics were accompanied by a decrease in epidermal cell number which was partly compensated for by an increase in epidermal cell area. Overall, the whole rosette leaf expansion rate was reduced by shading, whereas the total duration of rosette leaf expansion was unaffected. This was mainly due to the accumulation of the increases in the durations of expansion of each individual leaf which was associated with an increase in cell expansion. CONCLUSIONS: The dynamic analysis presented here reveals a new shade-adaptative response mediated via the control of area expansion at the cell, organ and whole plant levels.     

Effects of the fungal endophyte, Neotyphodium lolii, on net photosynthesis and growth rates of perennial ryegrass (Lolium perenne) are independent of In Planta endophyte concentration

• Background and Aims Neotyphodium lolii is a fungal endophyteof perennial ryegrass (Lolium perenne), improving grass fitnessthrough production of bioactive alkaloids. Neotyphodium speciescan also affect growth and physiology of their host grasses(family Poaceae, sub-family Pooideae), but little is known aboutthe mechanisms. This study examined the effect of N. lolii onnet photosynthesis (P_n) and growth rates in ryegrass genotypesdiffering in endophyte concentration in all leaf tissues. • Methods Plants from two ryegrass genotypes, Nui D andNui UIV, infected with N. lolii (E+) differing approx. 2-foldin endophyte concentration or uninfected clones thereof (E–)were grown in a controlled environment. For each genotype xendophyte treatment, plant growth rates were assessed as tilleringand leaf extension rates, and the light response of P_n, darkrespiration and transpiration measured in leaves of young (30–45d old) and old (>90 d old) plants with a single-chamber openinfrared gas-exchange system. • Key Results Neotyphodium lolii affected CO₂-limited ratesof P_n, which were approx. 17 % lower in E+ than E– plants(P < 0·05) in the young plants. Apparent photon yieldand dark respiration were unaffected by the endophyte (P >0·05). Neotyphodium lolii also decreased transpiration(P < 0·05), but only in complete darkness. There wereno endophyte effects on P_n in the old plants (P > 0·05).E+ plants grew faster immediately after replanting (P < 0·05),but had approx. 10 % lower growth rates during mid-log growth(P < 0·05) than E– plants, but there was noeffect on final plant biomass (P > 0·05). The endophyteeffects on P_n and growth tended to be more pronounced in NuiUIV, despite having a lower endophyte concentration than NuiD. • Conclusions Neotyphodium lolii affects CO₂ fixation,but not light interception and photochemistry of P_n. The impactof N. lolii on plant growth and photosynthesis is independentof endophyte concentration in the plant, suggesting that theendophyte mycelium is not simply an energy drain to the plant.However, the endophyte effects on P_n and plant growth are stronglydependent on the plant growth phase.     

Xylem structure and connectivity in grapevine (Vitis vinifera) shoots provides a passive mechanism for the spread of bacteria in grape plants

BACKGROUND AND AIMS: Bacterial leaf scorch occurring in a number of economically important plants is caused by the xylem-limited bacterium Xylella fastidiosa (Xf). In grapevine, Xf systemic infection causes Pierce's disease and is lethal. Traditional dogma is that Xf movement between vessels requires the digestion of inter-vessel pit membranes. However, Yersinia enterocolitica (Ye) (a bacterium found in animals) and fluorescent beads moved rapidly within grapevine xylem from stem into leaf lamina, suggesting open conduits consisting of long, branched xylem vessels for passive movement. This study builds on and expands previous observations on the nature of these conduits and how they affect Xf movement. METHODS: Air, latex paint and green fluorescence protein (GFP)-Xf were loaded into leaves and followed to confirm and identify these conduits. Leaf xylem anatomy was studied to determine the basis for the free and sometimes restricted movement of Ye, beads, air, paint and GFP-Xf into the lamina. KEY RESULTS: Reverse loading experiments demonstrated that long, branched xylem vessels occurred exclusively in primary xylem. They were observed in the stem for three internodes before diverging into mature leaves. However, this stem-leaf connection was an age-dependent character and was absent for the first 10-12 leaves basal to the apical meristem. Free movement in leaf blade xylem was cell-type specific with vessels facilitating movement in the body of the blade and tracheids near the leaf margin. Air, latex paint and GFP-Xf all moved about 50-60% of the leaf length. GFP-Xf was never observed close to the leaf margin. CONCLUSIONS: The open vessels of the primary xylem offered unimpeded long distance pathways bridging stem to leaves, possibly facilitating the spread of bacterial pathogens in planta. GFP-Xf never reached the leaf margins where scorching appeared, suggesting a signal targeting specific cells or a toxic build-up at hydathodes.     

Branch development controls leaf area dynamics in grapevine (Vitis vinifera) growing in drying soil

BACKGROUND AND AIMS: Soil water deficit is a major abiotic stress with severe consequences for the development, productivity and quality of crops. However, it is considered a positive factor in grapevine management (Vitis vinifera), as it has been shown to increase grape quality. The effects of soil water deficit on organogenesis, morphogenesis and gas exchange in the shoot were investigated. METHODS: Shoot organogenesis was analysed by distinguishing between the various steps in the development of the main axis and branches. Several experiments were carried out in pots, placed in a greenhouse or outside, in southern France. Soil water deficits of various intensities were imposed during vegetative development of the shoots of two cultivars ('Syrah' and 'Grenache N'). KEY RESULTS: All developmental processes were inhibited by soil water deficit, in an intensity-dependent manner, and sensitivity to water stress was process-dependent. Quantitative relationships with soil water were established for all processes. No difference was observed between the two cultivars for any criterion. The number of leaves on branches was particularly sensitive to soil water deficit, which rapidly and strongly reduced the rate of leaf appearance on developing branches. This response was not related to carbon availability, photosynthetic activity or the soluble sugar content of young expanding leaves. The potential number of branches was not a limiting factor for shoot development. CONCLUSIONS: The particularly high sensitivity to soil water deficit of leaf appearance on branches indicates that this process is a major determinant of the adaptation of plant leaf area to soil water deficit. The origin of this particular developmental response to soil water deficit is unclear, but it seems to be related to constitutive characteristics of branches rather than to competition for assimilates between axes differing in sink strength.     

Host recognition by a polyphagous lepidopteran (Helicoverpa armigera): primary host plants, host produced volatiles and neurosensory stimulation

Abstract An important question in the host‐finding behaviour of a polyphagous insect is whether the insect recognizes a suite or template of chemicals that are common to many plants? To answer this question, headspace volatiles of a subset of commonly used host plants (pigeon pea, tobacco, cotton and bean) and nonhost plants (lantana and oleander) of Helicoverpa armigera Hübner (Lepidoptera: Noctuidae) are screened by gas chromatography (GC) linked to a mated female H. armigera electroantennograph (EAG). In the present study, pigeon pea is postulated to be a primary host plant of the insect, for comparison of the EAG responses across the test plants. EAG responses for pigeon pea volatiles are also compared between females of different physiological status (virgin and mated females) and the sexes. Eight electrophysiologically active compounds in pigeon pea headspace are identified in relatively high concentrations using GC linked to mass spectrometry (GC‐MS). These comprised three green leaf volatiles [(2E)‐hexenal, (3Z)‐hexenylacetate and (3Z)‐hexenyl‐2‐methylbutyrate] and five monoterpenes (α‐pinene, β‐myrcene, limonene, E‐β‐ocimene and linalool). Other tested host plants have a smaller subset of these electrophysiologically active compounds and even the nonhost plants contain some of these compounds, all at relatively lower concentrations than pigeon pea. The physiological status or sex of the moths has no effect on the responses for these identified compounds. The present study demonstrates how some host plants can be primary targets for moths that are searching for hosts whereas the other host plants are incidental or secondary targets.     

International research on cassava photosynthesis, productivity, eco-physiology, and responses to environmental stresses in the tropics

The review sums up research conducted at CIAT within a multidiscipline effort revolving around a strategy for developing improved technologies to increase and sustain cassava productivity, as well as conserving natural resources in the various eco-edaphic zones where the crop is grown, with emphasis on stressful environments. Field research has elucidated several physiological plant mechanisms underlying potentially high productivity under favourable hot-humid environments in the tropics. Most notable is cassava inherent high capacity to assimilate carbon in near optimum environments that correlates with both biological productivity and root yield across a wide range of germplasm grown in diverse environments. Cassava leaves possess elevated activities of the C₄ phosphoenolpyruvate carboxylase (PEPC) that also correlate with leaf net photosynthetic rate (P _N) in field-grown plants, indicating the importance of selection for high P _N. Under certain conditions such leaves exhibit an interesting photosynthetic C₃-C₄ intermediate behaviour which may have important implications in future selection efforts. In addition to leaf P _N, yield is correlated with seasonal mean leaf area index (i.e. leaf area duration, LAD). Under prolonged water shortages in seasonally dry and semiarid zones, the crop, once established, tolerates stress and produces reasonably well compared to other food crops (e.g. in semiarid environments with less than 700 mm of annual rain, improved cultivars can yield over 3 t ha⁻¹ oven-dried storage roots). The underlying mechanisms for such tolerance include stomatal sensitivity to atmospheric and edaphic water deficits, coupled with deep rooting capacities that prevent severe leaf dehydration, i.e. stress avoidance mechanisms, and reduced leaf canopy with reasonable photosynthesis over the leaf life span. Another stress-mitigating plant trait is the capacity to recover from stress, once water is available, by forming new leaves with even higher P _N, compared to those in nonstressed crops. Under extended stress, reductions are larger in shoot biomass than in storage root, resulting in higher harvest indices. Cassava conserves water by slowly depleting available water from deep soil layers, leading to higher seasonal crop water-use and nutrient-use efficiencies. In dry environments LAD and resistance to pests and diseases are critical for sustainable yields. In semiarid zones the crop survives but requires a second wet cycle to achieve high yields and high dry matter contents in storage roots. Selection and breeding for early bulking and for medium/short-stemmed cultivars is advantageous under semiarid conditions. When grown in cooler zones such as in tropical high altitudes and in low-land sub-tropics, leaf P _N is greatly reduced and growth is slower. Thus, the crop requires longer period for a reasonable productivity. There is a need to select and breed for more cold-tolerant genotypes. Selection of parental materials for tolerance to water stress and infertile soils has resulted in breeding improved germplasm adapted to both favourable and stressful environments. An erratum to this article is available at .     

Morpho-anatomical and physiological leaf traits of two alpine herbs, Podophyllum hexandrum and Rheum emodi in the Western Himalaya under different irradiances

Morpho-anatomical leaf traits and photosynthetic activity of two alpine herbs, Podophyllum hexandrum (shade-tolerant) and Rheum emodi (light-requiring), were studied under field (PAR>2 000 μmol m⁻² s⁻¹) and greenhouse (PAR 500 μmol m⁻² s⁻¹) conditions. Mesophyll thickness, surface area of mesophyll cells facing intercellular spaces (S_mes), surface area of chloroplasts facing intercellular spaces (S_c), intercellular spaces of mesophyll cells (porosity), photon-saturated rate of photosynthesis per unit leaf area (P _Nmax), and ribulose-1,5-bisphosphate carboxylase/oxygenase activity decreased in the greenhouse with respect to the field and the decreases were significantly higher in R. emodi than in P. hexandrum. P. hexandrum had lower intercellular CO₂ concentration than R. emodi under both irradiances. The differences in acclimation of the two alpine herbs to low irradiance were due to their highly unlikely changes in leaf morphology, anatomy, and P _Nmax which indicated that the difference in radiant energy requirement related to leaf acclimation had greater impact under low than high irradiance.     

Acclimation to irradiance in seedlings of three tropical rain forest Garcinia species after simulated gap formation

We investigated the acclimation of seedlings of three tropical rain forest sub-canopy Garcinia species (G. xanthochymus, G. cowa, and G. bracteata) after transfer from 4.5 (LI) to 40 % (HI) sunlight and 12.5 (MI) sunlight to HI (LH₁ and LH₂ denoting transfer from LI to HI and MI to HI transfer, respectively). The changes of chlorophyll (Chl) fluorescence, net photosynthetic rate (P _N), dark respiration rate (R _D), Chl content per unit area (Chl_area), leaf mass per unit area (LMA), and seedling mortality were monitored over two months after transfer. These parameters together with leaf anatomy of transferred and control seedlings (kept in LI, MI, and HI) were also examined after two months. No seedlings died during the two months. F_v/F_m, P _N, and Chl_area of the transferred seedlings decreased in the first 3 to 12 d. LH₁ leaves showed larger reduction in F_v/F_m (>23 % vs. <16 %) and slower recovery of F_v/F_m than LH₂ leaves. P _N started to recover after about one week of I transfer and approached higher values in all G. cowa seedlings and G. xanthochymus LH₁ seedlings than those before the transfer. However, P _N of G. bracteata seedlings approached the values before transfer. The final P _N values in leaves of transferred G. xanthochymus and G. cowa seedlings approached that of leaves kept in HI, while the final P _N values of transferred leaves of G. bracteata were significantly lower than that of leaves grown under HI (p<0.05). R _D of G. xanthochymus LH₁ seedlings and all G. cowa seedlings increased and approached the value of the seedlings in HI. The final Chl_area of both G. xanthochymus and G. cowa approached the values before transfer, but that of G. bracteata did not recover to the level before transfer. The final Chl_area of all transferred seedlings was not significantly different from that of seedlings in HI except that G. cowa LH₁ seedlings had higher Chl_area than that in HI. LMA decreased within 2 d and then increased continuously until about 30 d and approached the value under HI. Spongy/palisade mesophyll ratio decreased after transfer because of the increase in palisade thickness. Leaf thickness did not change, so LMA increase of transferred seedlings was mainly due to the increase of leaf density. Thus the mature leaves under LI and MI of G. xanthochymus and G. cowa are able to acclimate to HI by leaf physiological and anatomical adjustment, while G. bracteata had limited ability to acclimate to HI.     

Relationship between δ13C and photosynthetic parameters and their responses to leaf nitrogen content in six broadleaf tree species

Stable carbon isotope composition (δ¹³C), net photosynthetic rate (P _N), actual quantum yield of photosystem 2 (PS2) electron transport (ΦPS2), nitrogen content (N_c), and photosynthetic nitrogen use efficiency (PNUE) in the leaves of six broadleaf tree species were determined under field environmental conditions. The six tree species were Magnolia liliflora Desr., M. grandiflora Linn., M. denudata Desr., Prunus mume (Sieb.) Sieb. et Zucc. cv. Meiren Men, P. mume (Sieb.) Sieb. et Zucc. f. alphandii (Carr.) Rehd., and P. persica (L.) Batsch. var. rubro-plena. The relationships among δ¹³C, Φ_PS2, P _N, and PNUE, as well as their responses to N_c in the six species were also studied. Both P _N and δ¹³C negatively correlated with N_c, but Φ_PS2 positively correlated with N_c. This indicated that with N_c increase, P _N and δ¹³C decreased, while Φ_PS2 increased. There were weak negative correlations between δ¹³C and PNUE, and strong negative correlations (p<0.01) between Φ_PS2 and PNUE. According to the variance analysis of parameters, there existed significant interspecific differences (p<0.001) of δ¹³C, P _N, Φ_PS2, PNUE, and N_c among the tree seedlings of the six tree species, which suggests that the potential photosynthetic capacities depend on plant species, irradiance, and water use capacity under field conditions.