The phytotoxin coronatine induces light-dependent reactive oxygen species in tomato seedlings

The phytotoxin coronatine (COR), which is produced by Pseudomonas syringae pv. tomato DC3000 (DC3000), has multiple roles in virulence that lead to chlorosis and a reduction in chlorophyll content. However, the physiological significance of COR-induced chlorosis in disease development is still largely unknown. Global expression analysis demonstrated that DC3000 and COR, but not the COR-defective mutant DB29, caused reduced expression of photosynthesis-related genes and result in a 1.5- to 2-fold reduction in maximum quantum efficiency of photosystem II (F(V)/F(M)). Tomato (Solanum lycopersicum) seedlings inoculated with DC3000 and incubated in a long daily photoperiod showed more necrosis than inoculated seedlings incubated in either dark or a short daily photoperiod. The accumulation of reactive oxygen species (ROS) was detected in cotyledons inoculated with either purified COR or DC3000 but not in tissues inoculated with DB29. Interestingly, COR-induced ROS accumulated only in light and was inhibited by 3-(3,4-dichlorophenyl)-1,1-dimethylurea and diphenylene iodonium, which function to inhibit electron transport from PSII. Furthermore, COR and DC3000 suppressed expression of the gene encoding the thylakoid Cu/Zn superoxide dismutase but not the cytosolic form of the same enzyme. In conclusion, these results demonstrate a role for COR-induced effects on photosynthetic machinery and ROS in modulating necrotic cell death during bacterial speck disease of tomato.     

Effects of Natural and Artificial Light in Arctic Latitudes on Long- and Short-day Plants as Revealed by Growth Analysis

The effects on growth and flowering of two short-day and twolong-day plants when grown under different conditions of illuminationare described. The plants fully investigated were Kalanchoeblossfeldiana and Xanthium pennsylvanicum and the annual varietiesof Hyoscyamus niger and Beta vulgaris. Wintex barley, Iberisumbellata, and tomato were also grown in some selected treatments.The conditions investigated comprised continuous full daylight(24 hours), full daylight for the whole of the daily photoperiodand full daylight for half the photoperiods, the other halfconsisting of either daylight reduced by shading or light fromincandescent lamps or fluorescent tubes (daylight-matching type),all of the same low intensity. Two lengths of photoperiod wereused for each species, one nearly optimal for flowering, theother closer to the critical day-length; and the order of thelow and high light treatments was varied. These factors werecombined factorially. Data were collected (or derived) for the following characteristics,though not always for all the species grown: dry weights, leafareas, heights, water contents, epidermal cell sizes, net assimilationrates, times to flowering, leaf-number increments until flowering,numbers of inflorescences, stomatal apertures, and leaf postures. Among other effects, the data revealed that in all four speciesinvestigated the adverse effects on over-all growth to be expectedfrom reduction of the daily photoperiod or of the intensityof illumination are in fact minimized. This compensation waseffected mainly by large increases in leaf areas, even thoughin all cases half the daily photoperiod consisted of full daylight.There are indications that increased efficiencies (net assimilationrates) may also have been involved. The leaf-area increasesappear to have been due to increased cell size rather than cellnumber and a close positive correlation with water content wasfound. The most striking among the effects on flowering was the failureof sugar-beet to bolt when half of its photoperiod (totals of20 and 14 hours) consisted of light from fluorescent lamps.The flowering of barley and Hyoscyamus was also delayed considerablyunder these conditions. The deficiency of red in the spectrumof the fluorescent light is believed to have been the cause.By contrast, the flowering of Iberis, a crucifer, was not affected.     

Adaptability of tomato and pepper leaves to changes in photoperiod: Effects on the composition and function of the thylakoid membrane

The adaptability of the thylakoid membrane to extended photoperiod (from natural to 24 h) was studied using a photoperiod-sensitive species ( Lycopersicon esculentum Mill. cv. Trend) and a non-photoperiod-sensitive species ( Capsicum annuum L. cv. Delphin). Our results have shown that thylakoid membranes of both species adapt to an extended photoperiod by increasing their photosystem II to photosystem I ratio (PSII/PSI) in order to provide a more balanced energy distribution between both photosystems to improve quantum yield. In tomato plants, these results correspond with a lower chlorophyll (Chl) a/b ratio, a decrease in Chl associated with PSI light-harvesting chlorophyll a/b protein complexes and with an increase in Chl associated with PSII light-harvesting chlorophyll a/b protein complexes. In spite of these changes, the electron transport capacity through PSII and PSI per unit of Chl and the light saturation point of PSII remained unchanged. The inability of tomato plants to use supplemental light for an extended photoperiod is not the result of photoinhibitory conditions. In pepper plants a significant increase in electron transport capacity and in the light saturation point of PSII was found. There was a significant increase in CO₂ assimilation when the light period was increased from 12 to 24 h. In contrast to tomato, pepper plants adapt to a 24-h photoperiod by increasing their carboxylation capacity which is accompanied by an increase in electron transport capacity and the light saturation point.     

NTRC and chloroplast-generated reactive oxygen species regulate Pseudomonas syringae pv. tomato disease development in tomato and arabidopsis

Coronatine (COR)-producing pathovars of Pseudomonas syringae, including pvs. tomato, maculicola, and glycinea, cause important diseases on tomato, crucifers, and soybean, respectively, and produce symptoms with necrotic lesions surrounded by chlorosis. The chlorosis is mainly attributed to COR. However, the significance of COR-induced chlorosis in localized lesion development and the molecular basis of disease-associated cell death is largely unknown. To identify host (chloroplast) genes that play a role in COR-mediated chlorosis, we used a forward genetics approach using Nicotiana benthamiana and virus-induced gene silencing and identified a gene which encodes 2-Cys peroxiredoxin (Prxs) that, when silenced, produced a spreading hypersensitive or necrosis-like phenotype instead of chlorosis after COR application in a COI1-dependent manner. Loss-of-function analysis of Prx and NADPH-dependent thioredoxin reductase C (NTRC), the central players of a chloroplast redox detoxification system, resulted in spreading accelerated P. syringae pv. tomato DC3000 disease-associated cell death with enhanced reactive oxygen species (ROS) accumulation in a COR-dependent manner in tomato and Arabidopsis. Consistent with these results, virulent strain DC3000 suppressed the expression of Prx and NTRC in Arabidopsis and tomato during pathogenesis. However, interestingly, authentic COR suppressed the expression of Prx and NTRC in tomato but not in Arabidopsis, suggesting that COR in conjunction with other effectors may modulate ROS and cell death in different host species. Taken together, these results indicated that NTRC or Prx function as a negative regulator of pathogen-induced cell death in the healthy tissues that surround the lesions, and COR-induced chloroplast-localized ROS play a role in enhancing the disease-associated cell death.     

Host plant, temperature, and photoperiod effects on ovipositional preference of Frankliniella occidentalis and Frankliniella fusca (Thysanoptera: Thripidae)

Host plant effects of tomato, Lycopersicon esculentum Mill., and chickweed, Stellaria media (L.) Vill., foliage infected and uninfected with Tomato spotted wilt virus (family Bunyaviridae, genus Tospovirus, TSWV) on the ovipositional preferences of western flower thrips, Frankliniella occidentalis (Pergande), and tobacco thrips, Frankliniella fusca (Hinds), were investigated for whole plants in the greenhouse. In addition, the preference for leaf disks from the same host plants was investigated under a range of temperatures, 15-30 degrees C at a photoperiod of 12:12 (L:D) h, and at three photoperiods, 6:18, 12:12, and 18:6, at 20 degrees C in no-choice and choice studies conducted in growth chambers. In a choice test, F. fusca oviposited significantly more eggs per whole plant foliage over a 7-d period than F. occidentalis by an average ratio of 3:1 over both tomato and chickweed. The optimum temperature for oviposition of F. occidentalis and F. fusca was 24.5 and 24.9 degrees C, respectively. Both species laid significantly more eggs under the longest daylight hours tested, 18:6, in the choice study. Temperature and photoperiod did not significantly interact in terms of thrips ovipositional preference. Ovipositional preference for chickweed or tomato foliage was different for each thrips species in the choice and no-choice tests. However, both thrips species laid significantly more eggs per square centimeter of leaf area in chickweed than in tomato in the whole plant choice test.     

Maize growth and ion absorption in Richter's solution at different flow rates

By comparing maize plants cultivated in standing nutrient solution with those from solutions flowing at different flow rates it has been established that absorption of nitrogen, potassium and especially of phosphorus was increased owing to the flow. There was likewise a relative rise in the distribution of nutrients to the overground parts of the plants. The content expressed per unit dry matter was increased only in the case of phosphorus; with nitrogen and potassium it was slightly lower than in the standing solution. Increasing amounts of iron were required under the conditions of flowing nutrient solutions to prevent chlorosis of the plants. The production of dry matter,NAR andRGR was also increased because of the flow. The flow considerably changed the habitus of the primary roots of the maize plants. The roots were longer, thinner and on the whole they contained relatively less dry matter (RWR). The lengthening of the roots is explained as a response to stimulation by the solution flow—the rheotropism.     

Changes of the diurnal and circadian (endogenous) mRNA oscillations of the chlorophyll a/b binding protein in tomato leaves during altered day/night (light/dark) regimes

Characteristic steady-state mRNA level oscillations were monitored for the chlorophyll a/b-binding (cab) protein in tomato plants grown under the natural day/night (light/dark) regime as well as under constant environmental conditions. This typical expression pattern was altered when plants were transferred to different light/dark regimes. For example, by shifting the light phase by six hours, a change of the time points of maximum and minimum of expression level was monitored, while the principal oscillation pattern remained the same. It appeared that the transition from dark to light is involved in determining the time points of minima and maxima of mRNA accumulation.After exposing tomato plants to an abnormal light/dark periodicity (e.g. six hours of alternating light/dark) an altered oscillation pattern was determined: within 24 hours two maxima of cab mRNA levels were detected. However, this entrained abnormal rhythm was not manifested at the molecular level and the circadian pattern reappeared under constant environmental conditions (e.g. darkness). This result favours the hypothesis that the oscillation pattern of the cab mRNA in tomato plants is not only endogenous but also hereditary.     

Physiological and biochemical responses resulting from nitrite accumulation in tomato (Lycopersicon esculentum Mill. cv. Ibiza F1)

The sensitivity of hydroponically cultivated tomato (Lycopersicon esculentum Mill. cv. Ibiza F1) submitted to nitrite treatments (0.25-10mM KNO(2)) for 7d was studied. Increasing nitrite levels in the culture medium led to several disruptions of tomato plants, reflected by reductions of both dry matter per plant, chlorophyll concentrations and the appearance of chlorosis symptoms at the leaf surface. This behaviour was accompanied by stimulation of nitrite, nitrate and ammonia ion accumulation, mainly in roots and old leaves. Higher proteolytic and gaiacol peroxidase (GPX, EC. 1.11.1.7) activities and malonyldialdehyde content were also noted. Protein content of the different plant organs was decreased by nitrite treatment. These physiological and biochemical parameters were chosen as they are stress indicators. Taken together, our data partly explain the harmful effects of nitrite ions, when excessive in the culture medium.     

The periodicity of primary increment formation in the otoliths of Arctic charr Salvelinus alpinus (L.)

The role of environmental factors in the regulation of sub-annual increment formation in fish otoliths appears to differ markedly between species. To examine the periodicity of primary increment formation in the otoliths of O + Arctic charr, Salvelinus alpinus (L.), and the effects of temperature, and photoperiod on their formation, fish were held under controlled environmental conditions. Primary growth increments were found in the otoliths of fish held at constant temperature (18° C) and at ambient temperature [fluctuating with a circadian and circannual rhythm (4–18° C)]. Consistent and significant disruptions in increment formation occurred however, in experimental groups subjected to rapid change from ambient photoperiod to a 6L: 6D photo-period for 96 h. Disruptions in increment formation were also observed immediately following transportation of fish between holding facilities and following disease treatment. The number of otolith increments formed in fish held on an ambient photoperiod regime, correlated closely with time elapsed in days since checkmark formation ( r = 0.989, P ≤0.001) in fish sampled sequentially over a period of 10 to 105 days. Thus we demonstrate that under conditions of ambient photoperiod, primary increments are formed daily.     

A single nucleotide change within a plant virus satellite RNA alters the host specificity of disease induction

Some RNA plant viruses contain satellite RNAs which are dependent upon their associated virus for replication and encapsidation. Some cucumber mosaic virus satellite RNAs induce chlorosis on any of several host plants, including either tobacco or tomato. The exchange of sequence domains between cDNA clones of chlorosis-inducing and non-pathogenic satellite RNAs delimited the chlorosis domain for both tobacco and tomato plants to the same region. Site-directed mutagenesis of one nucleotide (149) within this domain changed the host plant specificity for a chlorotic response to satellite RNA infection from tomato to tobacco. Within the chlorosis domain, three conserved nucleotides are either deleted or altered in all satellite RNAs that do not induce chlorosis. Deletion of one of these nucleotides (153) did not affect satellite RNA replication but rendered it non-pathogenic. Thus, two single nucleotides have been identified which play central roles in those interactions between the virus, its satellite RNA and the host plant, and together result in a specific disease state.     

Influence des héméropériodes très courtes sur la croissance de Lolium multiflorum,sa composition pigmentaire et l'ultrastructure chloroplastique

Summary We have described some characteristics of Lolium multiflorum cultivated under very short photoperiods (2 hours and 1 hour). The estimations of leaf growth were based on dry weight, surface measurements, and chlorophyll content. The pigment analyses were carried out by column chromatography; chloroplast ultrastructure was observed after chemical fixation.These measurements have permitted us to note a sharp drop in the growth curve of plants grown under different day-lengths: the limiting photoperiod lies between 1 hour and 2 hours of daily illumination.Pigment analyses and chloroplast ultrastructure observations show that there is a greater difference between plants cultivated under 1 hour and 2 hours of daily illumination than between plants cultivated under 2 hours and 12 hours.A decrease in day-length causes a deficit in the chlorophyll b content as well as a poor development of the grana.We have attempted to correlate these structural anomalies with the abnormal chlorophyll a/chlorophyll b ratio.     

Effects of Far-red Light on the Hormonal Control of Side Shoot Growth in the Tomato

Side shoot growth in young tomato plants was almost completelysuppressed by a 5 min period of far-red light immediately followinga 16 h photoperiod from fluorescent tubes, whereas plants givenan identical photoperiod but lacking the far-red treatment branchedprofusely. The influence of far-red light on the degree of sideshoot suppression and the correlated changes in the levels ofauxins, gibberellins, cytokinins and abscisic acid is presentedand discussed in relation to current hypotheses of correlativeinhibition. It is suggested that far-red light causes increasedauxin synthesis in the apex and young leaves, which in turninduces the formation of abscisic acid in or near the axillarybuds, and it is this hormone which inhibits bud outgrowth. Therole of cytokinins and gibberellins remains uncertain but theyprobably act in a sequential manner, the gibberellins promotingbud growth following cytokinin-mediated release from apicaldominance.     

Effect of nitrate nutrition on iron utilization by an Fe-efficient and an Fe-inefficient soybean cultivar

Summary The effects of increasing amount of nitrate nitrogen on the growth, dry matter production, ionic balance and the appearance of iron chlorosis in two soybean cultivars were studied. More nitrogen increased the dry matter production of the Fe-efficient cultivar Hawkeye and decreased that of the Fe-inefficient cultivar T-203. The plants of Hawkeye showed no iron deficiency symptoms whereas all the plants of the Fe-inefficient cultivar T-203 developed Fe deficiency after about one week following emergence. The degree of chlorosis in the cultivar T-203 was more pronounced as the amount of nitrate applied increased.Deceased     

The Growth and Development of the Wheat Apex: The Effects of Photoperiod on Spikelet Production and Sucrose Concentration in the Apex

Spring wheat (Triticum aestivum cv. Warimba) plants were grownin a controlled environment (20°C) in two photoperiods (8or 16 h). In the first instance, plants were maintained in eachof the photoperiods from germination onwards at the same irradiance(375 µE m^–2 s^–1). In the second case, allplants were grown in a long photoperiod until 4 days after double-ridgeinitiation when half the plants were transferred to a shortphotoperiod with double the irradiance (16 h photoperiod at225 or 8 h at 475 µE ^–2 s^–1). The rates of growth and development of the apices were promotedby the longer photoperiod in both experiments. Shoot dry weightgain was proportional to the total light energy received perday whereas the dry weight of the shoot apex increased withincreasing photoperiod even when the total daily irradiancewas constant. The principal soluble carbohydrate present in the shoot apexwas sucrose, although low concentrations of glucose and fructosewere found in the apices of long photoperiod plants late indevelopment. Sucrose concentration was invariably greater inthe slow-growing apices of short photoperiod plants, but roseto approach this level in the long photoperiod plants when theterminal spikelet had been initiated. Triticum aestivum, wheat, apex, spikelet initiation, photoperiod, flower initiation     

Photosynthate Partitioning into Starch in Soybean Leaves: I. Effects of Photoperiod versus Photosynthetic Period Duration

Photosynthesis, photosynthate partitioning into foliar starch, and translocation were investigated in soybean plants (Glycine max (L.) Merr. cv. Amsoy 71), grown under different photoperiods and photosynthetic periods to determine the controls of leaf starch accumulation. Starch accumulation rates in soybean leaves were inversely related to the length of the daily photosynthetic period under which the plants were grown. Photosynthetic period and not photoperiod per se appears to be the important factor. Plants grown in a 14-hour photosynthetic period partitioned approximately 60% of the daily foliar accumulation into starch whereas 7-hour plants partitioned about 90% of their daily foliar accumulation into starch. The difference in starch accumulation resulted from a change in photosynthate partitioning between starch and leaf residual dry weight. Residual dry weight is defined as leaf dry weight minus the weight of total nonstructural carbohydrates. Differences in photosynthate partitioning into starch were also associated with changes in photosynthetic and translocation rates, as well as with leaf and whole plant morphology. It is concluded that leaf starch accumulation is a programmed process and not simply the result of a limitation in translocation.     

Growth and development of cucumber Cucumis sativus L. in the prereproductive period under long photoperiods

When plants are grown in a greenhouse or in controlled environment growth rooms, prolonging the photoperiod, including towards continuous light, is one of the ways to increase plant productivity and energy savings. However, exposing some plant species to long photoperiods causes leaf injuries and growth reductions. We studied the effect of the photoperiod (8, 12, 16, 20, and 24 h) and photosynthetic photon flux density (60, 120, and 160 μmol/m² s PAR) on cucumber plants Cucumis sativus L. in a prereproductive period. It was shown that the response of the cucumber plants to a photoperiod over 20 h, including continuous light, depending on the plant age and light intensity, may include leaf movement or paraheliotropism, non-photochemical energy dissipation, and/or reversible photoinhibition of a reaction center of photosystem II, development of reversible chlorosis, reduction of a light-harvesting complex, and increase in the content of carotenoids. Reaction of immature and virginile plants to long photoperiods was different, which high-lights the need for experimental separation of the prereproductive period in terms of age states and consideration of this when developing crop production plan.     

Growth,water and nutrient status of plants in relation to patterns of variations in concentrations of dry matter and nutrient elements in base-to-top leaves

Summary Patterns of variations in dry matter concentrations in tomato plants reflected production and translocation of dry matter, implying the possibility of controlling and regulating growth and development of plants by use of dry matter concentration as a useful parameter.Dry matter concentrations, analogous to nutrient concentrations, varied depending on growth conditions, and on type, age and position of plant organs.Interpretation of patterns of variations in contents and concentrations of leaf dry matter in plants, grown under widely different conditions, agreed with the source/sink hypothesis.High water applications were associated with high dry matter concentrations in upper leaves of young pot plants with low sink capacity and with low dry matter concentrations in leaves of older, trough-grown plants with high sink capacity.Accumulation of dry matter in upper leaves of plants is suggested to be associated with development of secondary sinks and, accumulation of dry matter in lateral shoots is considered as a possible explanation of apical dominance.Water regime and transpiration influenced distribution of contents of dry and fresh matter and of absorbed nutrient elements. Redistribution was influenced by water regime.The term, distribution is in the following used in connection with not only absolute values (contents) but also relative values (concentrations).     

Lunar and daily spawning rhythms of Senegal sole Solea senegalensis

A periodicity of 29 days was observed in spawning rhythms in Senegal sole Solea senegalensis , with an acrophase around the last quarter and the new moon. In both spring and autumn, a very marked nocturnal spawning rhythm was registered, with spawning beginning after dusk and the acrophase occurring around 2300 hours. When the photoperiod was artificially extended (from 10L:14D to 14L:10D), S. senegalensis synchronized to the new photoperiod: spawning took place after the new 'dusk', the beginning gradually shifting from 2100 to 2300 hours and the acrophase from 2325 to 0032 hours. Under continuous light conditions, fish sustained rhythmicity for 2 days, with an acrophase at 2249 hours, which suggested the existence of an endogenous pacemaker controlling the daily spawning rhythm. These findings provided new insights for better understanding the reproductive physiology of this species and for optimizing the timing protocols of egg collection and larvae production in S. senegalensis aquaculture.     

Growth and Maintenance Respiration in Whole Plants, Tops, and Roots of Lolium multiflorum

Continuous measurements of CO₂-exchange were separately carried out on tops and roots of small swards of Lolium multiflorum grown in nutrient solution in growth chamber during 3–4 weeks. From these measurements, a daily carbon balance and accumulated dry matter could be established. The data were used to distinguish between two components of respiration, one proportional to growth or photosynthesis (growth respiration), the other proportional to plant dry weight (maintenance respiration). The separation of respiration in the two components was made by multiple regression analyses with daily photosynthesis or growth rate and accumulated dry matter as the independent variables. To ensure independency between the independent variables during the growth period, photosynthesis was varied by application of alternate three-day periods of high and low irradiance. From the two regression coefficients, the efficiency of converting assimilates into constructive growth (Y_G) and the maintenance coefficient (M) could be derived. Three experiments with varying length of photoperiod and dark period were carried out. The analyses were carried out for whole-plant respiration, respiration of tops and respiration of roots separately. Growth respiration for whole plants as well as for tops and for roots was lower — and hence the efficiencies higher — the longer the photoperiods were. Growth respiration and maintenance respiration were higher for roots than for tops. The high rate of root respiration may originate from release of HCO₃^? in exchange for NO₃^?. The parameters found can be utilized quantitatively in computer models of crop photosynthesis and respiration.     

Dry matter partitioning in a tomato plant: one common assimilate pool?

The influence of the distance (transport resistance) betweensource and sink on dry matter distribution between fruits andvegetative parts in tomato was studied. In two glasshouse experiments,a control treatment (single-shoot plants, no truss removal)was conducted, together with two double-shoot treatments: double-shootplants with no trusses removed from one shoot and all trussesremoved at anthesis from the other shoot (100–0) and double-shootplants with every second truss removed from both shoots (50–50).Plant growth and dry matter distribution was recorded by periodicaldestructive harvests, during a period of about 100 d after anthesisof the first truss. In experiment 2, plants were probably sink-limited.At the end of both experiments, 58–60% of dry matter wasin the fruits for control plants, whereas for both double-shoottreatments this was 43% (Experiment 1) or 38% (Experiment 2).Until 60–65 d after first flowering, vegetative growthof the individual shoots in both double-shoot treatments wasthe same. Results supported the assumption of one common assimilatepool and showed no significant influence of distance (transportresistance) between source and sink on dry matter partitioning. Key words: Allocation, distance, dry matter distribution, one assimilate pool, partitioning, transport resistance, tomato