SELECTIVITY AND PHYSIOLOGICAL INACTIVATION OF SOME HERBICIDES INHIBITING PHOTOSYNTHESIS

Summary. The effects of several herbicides on the rates of CO₂ uptake and transpiration of intact plants were measured. A simple technique is described which reveals whether photosynthesis-inhibiting herbicides are inactivated by plants. By this method, the inactivation of simazine by maize could be confirmed. Evidence is presented for physiological inactivation of monuron by Plantago lanceolata , of cycluron (OMU) and pyrazon by sugar beet, of N'-(4-butoxyphenyl)-NN-dimethylurea by of 5-bromo-6-methyl-3-phenyluracil by flax. Certain plant species showed different reactions to some other new herbicides. The effect of herbicides in the culture solution on CO₂ uptake by the leaves is probably largely determined by the rate of transpiration of the plants. Assuming that uptake in proportional to transpiration rate, estimates of the concentration of the herbicides inside the plant were made.
L'inactivation physiologique de quelques herbicides produisant une inhibition de la photosynthèse     

The effect of elevated CO2 levels on the symptom expression of Colletotrichum gloeosporioides on avocado fruits

Exposure of freshly harvested avocado fruits to different concentrations of CO₂ (11, 16 and 30%) for different lengths of time (4, 17 and 26 h) affected the decay development caused by Colletotrichum gloeosporioides. The delay in symptom development depended on the treatment given, the temperature regime of the fruit and time after harvest for treatment application. The most appropriate treatment was the application of 30% CO₂ for 24 h at a temperature of 20-25 C on the day of harvest. Treatment for shorter time periods, at lower temperatures or 50 h after harvest, resulted in a reduced response and, in some cases, enhanced symptom expression. Concentrations of 11 or 16% CO₂ were less effective than 30% CO₂ as the fruits became more mature. It is concluded that treatment of avocado fruits with high levels of CO: for a short period has the potential to provide an alternative means of controlling anthracnose.     

Cutting weeds with a CO2 laser

Stems of Chenopodium album . and Sinapis arvensis . and leaves of Lolium perenne . were cut with a CO₂ laser or with a pair of scissors. Treatments were carried out on greenhouse-grown pot plants at three different growth stages and at two heights. Plant dry matter was measured 2 to 5 weeks after treatment. The relationship between dry weight and laser energy was analysed using a non-linear dose–response regression model. The regression parameters differed significantly between the weed species. At all growth stages and heights S. arvensis was more difficult to cut with a CO₂ laser than C. album . When stems were cut below the meristems, 0.9 and 2.3 J mm⁻¹ of CO₂ laser energy dose was sufficient to reduce by 90% the biomass of C. album and S. arvensis respectively. Regrowth appeared when dicotyledonous plant stems were cut above meristems, indicating that it is important to cut close to the soil surface to obtain a significant effect. When cutting L. perenne plants with 2-true leaves at a height of 2 cm from the soil surface with a laser, the biomass decreased significantly compared with plants cut by scissors, indicating a delay in regrowth. This delay was not observed for the dicotyledonous plants nor for the other growth stages of L. perenne .     

Striga asiatica seed conditioning and 1-aminocyclopropane-1-carboxylate oxidase activity

Conditioned seeds of Striga asiatica (L.) Kuntze release ethylene, which elicits germination. We investigated the activity of 1-aminocyclopropane-1-carboxylate (ACC) oxidase and respiration during conditioning. Seeds incubated in vivo with ACC, the substrate for ACC oxidase, produced negligible ethylene at the beginning of conditioning or if they were dormant (i.e. would not germinate after conditioning and treatment with stimulant). Non-dormant seeds produced 3000 ηL of ethylene/600 seeds/24 h after 12 days of conditioning. In vitro ACC oxidase activity at day 0 of conditioning produced 500 ηL of ethylene/μg protein/h and 8000 ηL of ethylene/μg protein/h after 12 days of conditioning. Incubation of seeds in strigol before protein extraction did not enhance enzyme activity. Seeds released 4000 μL/L CO₂ in the first 24 h of conditioning, with the rate increasing to 15 000 μL/L/24 h on day 4 and then remaining roughly unchanged. Maximum in vitro activity of ACC oxidase required ACC, catalase, O₂, Fe²⁺, ascorbate and CO₂. In vivo activity of ACC oxidase required ACC and/or germination stimulant(s), suggesting that stimulants may be involved in providing substrates for the ACC oxidase. No difference was observed in the separation of extracted proteins, which suggests that ACC oxidase is activated during conditioning, perhaps as a result of changes in co-factor concentration. Application of these findings to Striga control is discussed.     

THE EFFECT OF SIMAZINE ON THE PHOTOFIXATION OF CO2 AND ON TRANSLOCATION OF ASSIMILATES IN NORWAY SPRUCE (PICEA ABIES)

Summary. A method of exposing seedlings of Norway spruce (Picea abies (L.) Karst.) to ¹⁴CO₂ is described. Within the 1st hr alter ¹⁴CO₂ exposure, no translocation of the ¹⁴C out of the treated branch could be observed. After a 24-hr period, however, the ¹⁴C in dormant seedlings had been translocated basipetally to part of the root system only, with no lateral diffusion of the ¹⁴C-compounds in the stem. About a week after exposure, both symplastic and apoplastic patterns of translocation had caused a more uniform distribution of ¹⁴C. In seedlings at active internode elongation, the translocation patterns were fundamentally identical to those in dormant seedlings, but the active shoot growth had led to a more uniform distribution of the ¹⁴C.
Simazine at 20 ppm had apparently stimulated both the photofixation of ¹⁴CO₂ and the rate of translocation of the ¹⁴C-assimilates. At 30 ppm, however, simazine had blocked the translocation of nutrients to the roots. On the other hand, the ¹⁴CO₂ uptake was not influenced. The simazine incubation had apparently no influence on the synthesis of cationic photosynthate.     

THE DECOMPOSITION OF UREA HERBICIDES BY BACILLUS SPHAERICUS, ISOLATED FROM SOIL

Summary. The decomposition of seven urea herbicides by Bacillus sphaericus was studied. Experiments with ¹⁴C-labelled monolinuron demonstrated that monolinuron was degraded by removal of CO₂ from the ureido portion of the molecule and leaving the /Kchloroaniline moiety. Tests with monuron, diuron, monolinuron, linuron, metobromuron, fluometuron and methabenzthiazuron indicated that B. sphaericus was only able to decompose the methoxy compounds monolinuron, linuron and metobromuron in this way. All compounds were identified by infrared spectroscopy. A pathway for degradation of the herbicides by B. sphaericus is proposed.     

Growth and photosynthesis of four invasive aquatic plant species in Europe

Crassula helmsii , Hydrocotyle ranunculoides , Ludwigia grandiflora and Myriophyllum aquaticum are four well known invasive aquatic plants in European waters. In this study, plant growth at different nutrient availabilities, regeneration capacity and photosynthesis were investigated. Results show high relative growth rates (RGR) of the species of up to 0.132 ± 0.008 g g⁻¹ dry weight (dw) day⁻¹ ( H. ranunculoides ) and a significant increase in RGR with increasing nutrient availability. All species show a high regeneration capacity and the ability to form new shoots from single nodes, even though it differs between the species. Ludwigia grandiflora and M. aquaticum also show regeneration from single leaves. Species differed in maximal amounts, and in temperature and light optima of net assimilation rates: H. ranunculoides leaves reach maximum photosynthetic rates of up to 3500 μmol CO₂ × h⁻¹ g⁻¹ dw, L. grandiflora (leaves) up to 2200 μmol CO₂ × h⁻¹ g⁻¹ dw, M. aquaticum (shoots) 400   μmol CO₂ × h⁻¹ g⁻¹ dw and C. helmsii (shoots) up to 200 μmol CO₂ × h⁻¹ g⁻¹ dw. Hydrocotyle ranunculoides , L. grandiflora and M. aquaticum preferred high light intensity and high temperatures, whilst C. helmsii was negatively affected by intense sunlight. Summarising, it can be assumed that at least H. ranunculoides , L. grandiflora and M. aquaticum can grow well under current and likely future central European climate conditions.     

AN AUTORADIOGRAPHIC STUDY OF THE MOVEMENT OF 14C-LABELLED ASSIMILATES IN THE DEVELOPING WHEAT PLANT

Summary. ¹⁴CO₂ together with autoradiographic techniques was used to stuy the movement of labelled carbon within wheat plants at different stages in their development. Movement of ¹⁴C about the plain was found to be extensive in the early stages, gradually bccoming more localized until at a stage after ear emergence there was little transference of labelled carbon between tillers within a 24-hour period.
In the early stages of development ¹⁴C from leaves on the main shoot was transported throughout the plant but accumulated in greatest amounts in the meristematic regions. Assimilates produced by the leaves of newly formed tillers were distributed mainly to the meristematic regions of those tillers although labelled carbon was moved in into the rest of the plant. In the later stages of development of each tiller the labelled assimilate was restricted to the tiller itself and to any very young tillers associated with it. After car emergence the pattern of distribution of the labelled carbon from the youngest and second youngest leaves was found to differ. Movement was predominantly towards the ear from the flag leaf and towards the root system from the leaf below the flag.
It was noticed that mature leaves accumulated some labelled material and the possibility that this occurred via the transpiration stream following migration from phloem to xylem, as is the case with some of the case with some of the translocated herbicides, was investigated, but no indication of xylem transference was obtained.
The possible application of these studies to investigations into the relationship between the movement natural assimilates and of translocated herbicides is discussed.     

TRANSLOCATION OF 14C-ASSIMILATES AND 3-AMINO-1,2,4-TRIAZOLE AND ITS METABOLITES IN AGROPYRON REPENS

Summary. Translocation of 3-amino-1,2,4-triazole-5-¹⁴C (¹⁴C-aminotriazole) was compared to that of ¹⁴C-assimilates in couch grass ( Agropyron repens (L.) Beauv.) at three different growth stages.
Assimilates of ¹⁴CO₂ were translocated from the treated shoot to other shoots and rhizomes at the 2–3-leaf and 3–4-leaf stages of development. Much less labelled material was translocated into untreated shoots at the 5-leaf stage. More ¹⁴C-assimilates were translocated to the roots than to untreated shoots at all developmental stages. The translocation patterns of ¹⁴C-aminotriazole and ¹⁴C-assimilates were similar.
Two metabolites, A and B, were formed from ¹⁴C-aminotriazole, which chromatographed identically to previously described metabolites in Cirsium arvense (L.) Scop. A was further metabolized into B. Labelled aminotriazole and its two metabolites were translocated throughout the plants. Metabolite A was phytotoxic when concentrated and re-applied to couch grass, but its properties were not those of Unknown II from Cirsium arvense. They were the same as those of Unknown III. Whether or not metabolite A and Unknown III are identical was not established.
Migration des substances assimilables marquées au ¹⁴ C, du 3 amino-1,2,4-triazole et de ses métabolites chez Agropyron repens     

Growth- and leaf-temperature effects on photosynthesis of sweet orange seedlings infected with Xylella fastidiosa

The effects of growth and leaf temperature on photosynthesis were evaluated in sweet orange seedlings ( Citrus sinensis cv. Pera) infected with Xylella fastidiosa (the bacterium that causes citrus variegated chlorosis, CVC). Measurements of leaf gas exchange and chlorophyll  a fluorescence were taken at leaf temperatures of 25, 30, 35 and 40°C in healthy and infected (without visible symptoms) seedlings submitted to two temperature regimes (25/20 or 35/20°C, day/night), not simultaneously. The CO₂ assimilation rates ( A ) and stomatal conductance ( g _s) were higher in healthy plants in both temperature regimes. Values for A and g _s of infected and healthy plants were higher in the 35/20°C regime, decreasing with leaf temperature increase. In addition, differences between healthy and infected plants were higher at 35/20°C, while no differences in chlorophyll  a fluorescence parameters were observed except for potential quantum efficiency of photosystem II, which was higher in infected plants. Low A values in infected plants were caused by low g _s and probably by biochemical damage to photosynthesis. The high alternative electron sink of infected plants was another effect of reduced A . Both high growth and high leaf temperatures increased differences in A between healthy and infected plants. Therefore this feature may be partially responsible for lower growth and/or productivity of CVC-affected plants in regions with high air temperature.     

AN AUTORADIOGRAPHIC STUDY OF THE DISTRIBUTION OF 14C LABELLED ASSIMILATES AT DIFFERENT STAGES OF DEVELOPMENT OF OXALIS LATIFOLIA H.B.K. AND O. PES-CAPRAE L.

Summary. An autoradiographic study of the distribution of ¹⁴C-labelled assimilates, derived from ¹⁴CO₂ fed to the aerial parts, was made at ten different developmental stages of Oxalis latifolia H.B.K. and Oxalis pes-caprae L. Changing patterns of distribution of labelled assimilates to the underground parts of both species were revealed. These patterns are discussed in the context of the timing of control by defoliation and by foliageapplied herbicides.
Etude autoradiographique de la distribution des métabolites marqués an ¹⁴C à différents stades de développement de l'Oxalis latifolia H.B.K. et de l' O. pes-caprae L.     

Weed shoot morphology effects on competitiveness for light in direct-seeded rice

The effects of weed shoot morphology on competitiveness for light in rice ( Oryza sativa L.) have not been well described quantitatively and are difficult to study empirically. A rice:weed model was used to analyse the effects of weed leaf area densities (LAD; m² m^–3), leaf angles (as leaf light extinction coefficients, k _leaf) and maximum heights ( H _m, m) on growth and competition with rice. Weed morphologies were hypothetical but empirically based, as follows: LADs were skewed to the bottom or conical, k _leaf values varied from 0.2 (erectophile) to 0.8 (planophile), and H _m values were 0.5 H _R, 1 H _R and 1.25 H _R, where H _R was rice maximum height. Other parameters were equal to those of rice. Growth and competitiveness were evaluated using mature seed dry weights (g m^–2). Short weeds and weeds with conical LADs were weakly competitive, regardless of other traits. For other weed types, interference with rice was positively related to H _m, LAD skewness and more planophile leaves. All three traits were critical determinants of weed interference but no single morphological trait guaranteed competitiveness. All else being equal, weeds with highly skewed LADs produced the most seed dry weight. Planophile leaves were particularly beneficial for short weeds, giving over five times more seed dry weight than erectophile leaves.     

EFFECT OF PYRAZON ON PHOTOSYNTHESIS OF VARIOUS PLANT SPECIES

Summary. Studies were conducted to determine the influence of pyrazon (5-amino-4-chloio-2-phenyl-3(2H)-pyridazinone) on photosynthesis. A strong inhibiting effect on O₂ evolution from leaf discs of several species was recorded. For most species a concentration of 5 × 10⁻⁵ M caused a reduction of about 50% in the rate of photosynthesis. Photosynthesis of sugar beet, highly resbtant to pyrazon, was reduced to 50% by a concentration of 7–5 × 10⁻⁵ M. Since these differences cannot account for the great divergence in susceptibility to pyrazon, it is suggested that the selectivity of this herbicide results from degradation and differential accumulation in the foliage.
Pyrazon was phytotoxic to plants of tomato and pigweed (Amaranthus retrofiexus L.) kept in the light but not when kept in the dark, after foliage application. The conclusion that pyrazon inhibits the mechanism of photosynthesis and causes starvation for carbohydrates, was strengthened by the fact that external feeding of glucose overcame the toxicity of the compound in tomato plants. These results and a possible mode of selectivity are discussed.     

Inheritance of tolerance to metribuzin in durum wheat

The inheritance of tolerance to the herbicide metribuzin was studied in two durum wheat cultivars, one of which, 'Anton', is resistant and one, 'Nita', susceptible. Parents, F₁, F₂ and F₃ of the crosses 'Anton' × 'Nita' and 'Nita' × 'Anton' were tested for herbicide response. The character evaluated was the increase in weight of plants treated with the herbicide. As there were no significant differences between progenies of reciprocal crosses, cytoplasm was not involved in tolerance. Tolerance was semi-dominant with means values of F₁, F₂ and F₃ progenies in the range of mid-parent. The heritability of this trait estimated by regression of the average of progeny F₃ in their parent F₂ had a value of 0.23 ± 0.063 and a value of 0.52 ± 0.150 estimated by the relation V _G/ V _P for full-sib F₂ families.     

Response of interspecific Brassica juncea/Brassica rapa hybrids and their advanced progenies to Albugo candida (white blister)

Transfer of factors for resistance to white blister disease caused by Albugo candida between Brassica species involving two genotypes each of B. juncea and B. rapa was studied in hybrids. More hybrids were obtained by in vivo than in vitro techniques, although an in vitro phase was a prerequisite for the establishment of in vivo hybrids. Hybrids were identified by PCR-based inter-simple sequence repeat (ISSR) markers with both male and female species-specific bands being identified. There was a positive correlation between disease severity and number of days after sowing ( r  > 0·93), the highest being towards pod formation and plant maturity at 110 days after sowing. The plants from F₂ and BC₁ progeny showed higher resistance to A. candida than either of the parents. Plants of B. juncea and B. rapa with high field resistance (disease index < 1·0) were selected from BC₂ and F₂BC₁ generations. The frequency of plants classified as resistant in BC₂ progeny ranged from 4·5 to 39·0% in cross-combinations involving B. juncea genotypes as female parent, compared with 100% in the reciprocal cross involving B. rapa as female parent.     

Physiological alterations in pepper during wilt induced by Phytophthora capsici and soil water deficit

Capsicum annuum cv. Morrón inoculated at the collar with Phytophthora capsici developed cortical stem necrosis and severe wilting soon after infection. Water potential, specific conductivity, ethylene evolution, CO₂ and water vapour exchange rates and chlorophyll content were monitored during symptom development. Well watered and non-inoculated water-stressed plants were used as controls. Necrosis developed upwards from the collar 4 days after inoculation coupled with a 80% reduction in hydraulic conductivity. Leaf water potential progressively decreased from -0.17 to -2.41 MPa in 2 weeks. The rate of ethylene evolution was significantly higher in infected than in non-inoculated stressed plants for similar water potentials and originated in the necrotic segment of disease stem tissue. Net photosynthesis and leaf conductance markedly decreased (74%) 4 days after infection, coupled to a burst in dark respiration (1.5 times) but were not associated with water stress alone. This suggests that the pathogen reduced photosynthesis initially through stomatal closure which was not directly mediated by water stress. The possible implication of ethylene in impaired stem conductivity and altered gas exchange of infected plants is discussed.     

Effects of Ascochyta blight (Mycosphaerella pinodes) on the photosynthesizing leaf area and the photosynthetic efficiency of the green leaf area of dried-pea (Pisum sativum)

The effects of Ascochyta blight (caused by Mycosphaerella pinodes ) on the net photosynthetic rate of glasshouse-grown dried pea were studied on foliar discs and whole plants in pots showing different disease intensities, assessed visually using a disease scale (scores 0 to 5) or by an estimation of the fraction of leaf area with necrosis using an image analyser. The photosynthetic rate of foliar discs was measured using a leaf disc electrode; for plants it was assessed by CO₂ exchange rate measurements in a closed chamber. A reduction in the net photosynthetic rate of diseased plants was correlated with an increase in disease score. A 53–56% reduction was found in foliar discs with a mean disease score of 3. On whole plants, mean scores of 3.5 and 4.0 were associated with reductions of 36% and 98% respectively. The disease also induced a decrease in the photosynthetic efficiency of the non-necrotic leaf area. The photosynthetic rate was zero when 30–40% of foliar area was necrotic on foliar discs and on whole plants. A function was derived from the foliar disc data that described the dependence of photosynthetic rate on the fraction of leaf area without necrosis. The model is discussed with reference to data obtained from entire plants and other pathosystems.     

INFLUENCE OF SOIL PROPERTIES ON PHYTOTOXICITY OF 4–AMlNO-3,5,6–TRICHLOROPICOLINIC ACID (PICLORAM)

Summary. Effective dosages of picloram (4–amino-3,5,6–trichloropicolinic acid) required to reduce fresh weight of sunflowers ( Helianthus annuus L. var. Menonite) were determined for seven Saskatchewan soils under controlled environmental conditions. The relationships between ED₅₀ value and clay content, organic matter content and cation exchange capacity were evaluated by correlation and regression analysis for possible usefulness in predicting dose requirements. ED₅₀ values were also determined for Weyburn loam and in culture solutions adjusted to various pH levels.
There Was no significant correlation between ED₅₀ values of picloram and soil clay content or cation exchange capacity. ED₅₀ values were highly correlated with soil organic matter content, and they increased as the soil pH was lowered or raised from pH 6.5. The increase in ED₅₀ values in the acidic range was attributed to adsorption of the unionized molecules of picloram on the organic matter in the soil. The increase in ED₅₀ values in the alkaline range may be mainly due to reduced uptake of the ionized acid by plant roots.     

Genetics of resistance to cotton leaf curl disease in Gossypium hirsutum

Twenty-two cotton varieties were screened for resistance to cotton leaf curl disease (CLCuD), a disease of viral origin, using three procedures: field evaluation, whitefly transmission assay and graft inoculation. Viral infection of cotton varieties was determined by visual symptom assessment as well as dot-blot and multiplex PCR diagnostic techniques. Crosses were made between the most susceptible variety (S-12) and highly resistant varieties (CP-15/2, LRA-5166 and CIM-443). All F₁ plants of these crosses were resistant, showing dominant expression of the resistance as well as the absence of extrachromosomal inheritance. The F₂ plants of the crosses CP-15/2 × S12, LRA-5166 × S-12 and CIM-443 × S12 exhibited a ratio of 13 resistant (symptomless) to three susceptible (with symptoms). Screening of the F₂ generation for virus infection by multiplex PCR further subdivided the resistant class into those exhibiting a high level of resistance (HR; PCR-negative) and those exhibiting resistance (R; symptomless, yet showing virus replication by PCR analysis). Hence, the final ratio was 3:10:3 (HR:resistant:susceptible). The F₃ progeny of susceptible F₂ plants segregated for resistance, indicating the probable presence of a suppressor gene ( S ). These findings are consistent with three genes being involved in G. hirsutum resistance to CLCuD, two for resistance ( R _1CLCuDhir and R _2CLCuDhir ) and a suppressor of resistance ( S _CLCuDhir ).     

UPTAKE, TRANSLOCATION AND METABOLISM OF SIMAZINE IN NORWAY SPRUCE (PICEA ABIES)

Summary. Uptake, distribution and fate simazine (2-chioro-4,6-biserthylamino- s -triazine) in 4-year-old seedlings of Norway spruce ( Picea abies ) are discussed. Simazine was found to be readily absorbed and distributed in the spruce seedlings. A positive relationship between the amount of simazine absorbed and the growth intensities of the plants indicates an active mechanism of uptake. A fixation of simazine in roots and sterm was observed.
Simazine was degraded by roots and stem to hydroxysimazine and two other unknown metabolites. No simazine and only metabolites could be traced in the needles, indicating a very rapid rate of metabolism in the root system. The glucose derivative of benzoxazinone was extracted from all parts of the spruce seedlings and is probably responsible for the hydrolysis of simazine.
The results indicate a very slow degradation of the metabolites to CO₂ as one of the final products.