Les herbicides inhibateurs du photosystème II, effets sur les communautés algales et leur dynamique

Nous présentons une revue bibliographique à propos des effets des herbicides inhibiteurs du Photosystème II (PS II) sur les communautés algales. Ces herbicides sont abondamment utilisés dans les pratiques phytosanitaires. Ils sont susceptibles de contaminer les milieux aquatiques et, étant donné leur mode d'action inhibitrice de la photosynthèse, ils peuvent agir directement sur les algues. De nombreuses études ont été réalisées afin d'évaluer l'impact des contaminations par ces herbicides sur les microphytes, en particulier leur effet sur la croissance et la physiologie de certaines algues (monocultures en laboratoire). D'autres études expérimentales et quelques rares in situ, ont porté sur l'impact des herbicides inhibiteurs de la photosynthèse sur la structure des peuplements algaux. Certaines tendances ont pu être ainsi dégagées quant à la sensibilité et la résistance aux herbicides des différentes espèces étudiées soit isolément, soit au sein des peuplements. Les herbicides inhibiteurs du PS II perturbent effectivement la structure des peuplements phytoplanctoniques de façon plus ou moins marquée. L'impact des herbicides sur les algues est variable selon la structure des peuplements (liée aux successions) et les paramètres environnementaux, notamment liés à la saison. Nous devons donc développer nos connaissances à propos des interactions entre toxiques et facteurs environnementaux sur des pas de temps correspondant non seulement aux rythmes des contaminations mais aussi aux rythmes des succesions alagles, car ces interactions sont susceptibles de réduire ou d'amplifier les conséquences d'une pollution par ces toxiques dans les milieux aquatiques.The aim of this paper is to present a review about the impact of Photo system II (PS II) inhibitors on algae communities. A brief discussion of the use in agriculture, the different chemical families, the photosynthetic inhibition effect and the occurrence of these compounds in aquatics systems is followed by the presentation of the impacts these herbicides have on algae. Many studies investigate the effects of PS II inhibitors on algae growth and physiology. The response to pollutants were studied by monitoring changes in terms of different parameters : chlorophyll fluorescence induction usually increases with PS II inhibitors. Concentration of pigments decreases with PS II inhibitors, but increases sometimes with low contaminations of these toxicants (it is probably an homeostasis effect). Pigment ratio can change with herbicide exposure. Primary production (measured by 14C incorporation or dissolved O2) usually decreases with PS II inhibitors. But, the " excretion " of dissolved organic compounds may increase with PS II inhibitors. These herbicides may alter or change cell morphology of algae. In consequence algae growth is inhibited by PS II inhibitors exposure. But growth inhibition varies, depending on each species' (and strain) sensibility or resistance to each herbicide. By this way, PS II inhibitors can affect the algae community structure. In consequence, herbicides exert a selection pressure when the exposure reaches a certain level, and this for a sufficient period of time. Since organisms vary in their resistance to toxicants, the selection pressure will exclude the sensitive ones which will be replaced by resistant ones. Sometimes, responses to pollutants which are measured by global changes in biomass, pigments, dissolved O2... can recover after a lagtime. This apparent ability to recover from effects of herbicides can be explained by the following selection effect : resistant species are indirectly stimulated and develop in the contaminated environment. The result is an algae community which has an increased resistance to these toxicants as compared to a community which has not been affected by the toxicants. This difference in resistance, between the unselected and the selected communities, may be detected by comparison of results from short term physiological tests performed with the respective community and by comparison of each community structure (taxonomy). This methodological approach called Pollution-Induced Community Tolerance (PICT) is of a great interest as a biological marker of specific pollution in aquatic systems. Indirect effects of algae response to PS II inhibitors occur in the polluted ecosystem according to changes of the physicochemical conditions (decrease of dissolved O2 concentrations and pH, increase of dissolved organic and inorganic matters ...). Furthermore, the impact of herbicides on algae communities varies, depending on the community's species composition (depending on successions) and on environmental factors such as interspecific interactions and physicochemical parameters (depending on seasonal changes). Interspecific interactions implies competition for the limiting nutrients among algae and allelopathic interactions among algae, as well as interactions between algae and other trophic levels (microbial loop, grazing pressure ...). These interactions between herbicides and environmental factors may reduce or emphasize the consequences of such a pollution in aquatic systems. Seasonal change of algae communities species composition (algae successions) occurs as a response to changing environmental factors by the way of interspecific interactions and physicochemical parameters. Therefore, algae succession is affected by the herbicide destructuration of the algae communities. At the opposite, interactions and successions may affect the response of algae communities to the toxic. In this sense, herbicides act as a supplementary factor of disturbance in algae successions. Structural changes, induced by these herbicides, are usually accompanied by the attributes which are typical of an early successional stage. But, according to the " Intermediate Disturbance Hypothesis " the species richness should be maximal at intermediate intensities of herbicide contamination and at intermeadite frequencies of contaminations. The question is to compare the algae successions rythms and the frequencies of herbicides contaminations. Then, the time factor (or the persisting quality, which is difficult to assess in experimental studies) has to be taken in account in monitoring aquatic polluted systems, because of the seasonal variability in the response of algae to PS II inhibitors as well as the seasonal variability of water bodies' contamination by these herbicides through watersheds. Moreover, usually, low values of herbicides' concentrations occur in aquatic environments but are persistant. It results that aquatic organisms are exposed during long periods of time, meaning that indirect effects, via interactions between herbicides and environmental factors, may be emphazised. To further investigate these interactions and the herbicide persistance in aquatic systems, we have to develop experimental studies. This approach, however, must be complemented with in situ studies monitored with a timing of investigation relative to the natural population fluctuations and " pulses " of herbicides in these systems. Investigation must take place on various aquatic ecosystems. If a greater effort is given to monitor natural systems, for both herbicides and herbicides-induced effects, this will provide greater confidence in future predictions regarding the safety of PS II inhibitors in aquatic environments

Role of isospin dependent mean field in pion production in heavy ion reactions

The importance of a isospin dependent nuclear mean field (IDMF) in regard to the pion production mechanism is studied for the reaction $Au+Au$ at 1 GeV/nucleon using the Quantum Molecular Dynamics (QMD) model. In particular, the effect of the IDMF on pion spectra and the charged pion ratio are analyzed. It is found that the inclusion of a IDMF considerably suppresses the low$-p_t$ pions, thus, leading to a better agreement with the data on pion spectra. Moreover, the rapidity distribution of the charged pion ratio appears to be sensitive to the isospin dependence of the nuclear mean field.Comment: 16 pages, using RevTex, 6 PS-Figure

Influence of Impact Parameter on Thermal Description of Relativistic Heavy Ion Collisions at GSI/SIS

Attention is drawn to the role played by the size of the system in the thermodynamic analysis of particle yields in relativistic heavy ion collisions at SIS energies. This manifests itself in the non-linear dependence of K+ and K- yields in $AA$ collisions at 1 -- 2 A.GeV on the number of participants. It is shown that this dependence can be quantitatively well described in terms of a thermal model with a canonical strangeness conservation. The measured particle multiplicity ratios (pi+/p, pi-/pi+, d/p, K+/pi+ and K+/K- but not eta/pi0) in central Au-Au and Ni-Ni collisions at 0.8 -- 2.0 A.GeV are also explained in the context of a thermal model with a common freeze-out temperature and chemical potential. Including the concept of collective flow a consistent picture of particle energy distributions is derived with the flow velocity being strongly impact-parameter dependent.Comment: revtex, 20 figure

K^+ production in baryon-baryon and heavy-ion collisions

Kaon production cross sections in nucleon-nucleon, nucleon-delta and delta-delta interactions are studied in a boson exchange model. For the latter two interactions, the exchanged pion can be on-mass shell, only contributions due to a virtual pion are included via the Peierls method by taking into account the finite delta width. With these cross sections and also those for pion-baryon interactions, subthreshold kaon production from heavy ion collisions is studied in the relativistic transport model.Comment: to appear in Phys. Rev.

Neutral Pions and Eta Mesons as Probes of the Hadronic Fireball in Nucleus-Nucleus Collisions around 1A GeV

Chemical and thermal freeze-out of the hadronic fireball formed in symmetric collisions of light, intermediate-mass, and heavy nuclei at beam energies between 0.8A GeV and 2.0A GeV are discussed in terms of an equilibrated, isospin-symmetric ideal hadron gas with grand-canonical baryon-number conservation. For each collision system the baryochemical potential mu_B and the chemical freeze-out temperature T_c are deduced from the inclusive neutral pion and eta yields which are augmented by interpolated data on deuteron production. With increasing beam energy mu_B drops from 800 MeV to 650 MeV, while T_c rises from 55 MeV to 90 MeV. For given beam energy mu_B grows with system size, whereas T_c remains constant. The centrality dependence of the freeze-out parameters is weak as exemplified by the system Au+Au at 0.8A GeV. For the highest beam energies the fraction of nucleons excited to resonance states reaches freeze-out values of nearly 15 %, suggesting resonance densities close to normal nuclear density at maximum compression. In contrast to the particle yields, which convey the status at chemical freeze-out, the shapes of the related transverse-mass spectra do reflect thermal freeze-out. The observed thermal freeze-out temperatures T_th are equal to or slightly lower than T_c, indicative of nearly simultaneous chemical and thermal freeze-out.Comment: 42 pages, 12 figure

Systematics of pion emission in heavy ion collisions in the 1A GeV regime

Using the large acceptance apparatus FOPI, we study pion emission in the reactions (energies in GeV/nucleon are given in parentheses): 40Ca+40Ca (0.4, 0.6, 0.8, 1.0, 1.5, 1.93), 96Ru+96Ru (0.4, 1.0, 1.5), 96Zr+96Zr (0.4, 1.0, 1.5), 197Au+197Au (0.4, 0.6, 0.8, 1.0, 1.2, 1.5). The observables include longitudinal and transverse rapidity distributions and stopping, polar anisotropies, pion multiplicities, transverse momentum spectra, ratios for positively and negatively charged pions of average transverse momenta and of yields, directed flow, elliptic flow. The data are compared to earlier data where possible and to transport model simulations.Comment: 56 pages,42 figures; to be published in Nuclear Physics

Isospin dependence of relative yields of K+K^+ and K0K^0 mesons at 1.528 AGeV

Results on $K^+$ and $K^0$ meson production in $^{96}_{44}$Ru + $^{96}_{44}$Ru and $^{96}_{40}$Zr + $^{96}_{40}$Zr collisions at a beam kinetic energy of 1.528$A$ GeV, measured with the FOPI detector at GSI-Darmstadt, are investigated as a possible probe of isospin effects in high density nuclear matter. The measured double ratio ($K^+/K^0$)$_{Ru}$/($K^+/K^0$)$_{Zr}$ is compared to the predictions of a thermal model and a Relativistic Mean Field transport model using two different collision scenarios and under different assumptions on the stiffness of the symmetry energy. We find a good agreement with the thermal model prediction and the assumption of a soft symmetry energy for infinite nuclear matter while more realistic transport simulations of the collisions show a similar agreement with the data but also exhibit a reduced sensitivity to the symmetry term.Comment: 5 pages, 3 figures. accepted for publication in Phys. Rev.

Transition from in-plane to out-of-plane azimuthal enhancement in Au+Au collisions

The incident energy at which the azimuthal distributions in semi-central heavy ion collisions change from in-plane to out-of-plane enhancement, E_tran, is studied as a function of mass of emitted particles, their transverse momentum and centrality for Au+Au collisions. The analysis is performed in a reference frame rotated with the sidewards flow angle, Theta_flow, relative to the beam axis. A systematic decrease of E_tran as function of mass of the reaction products, their transverse momentum and collision centrality is evidenced. The predictions of a microscopic transport model (IQMD) are compared with the experimental results.Comment: 32 pages, Latex, 22 eps figures, accepted for publication in Nucl. Phys.

Measurement of K∗(892)0K^*(892)^0 and K0K^0 mesons in Al+Al collisions at 1.9AA GeV

New measurement of sub-threshold $K^*(892)^0$ and $K^0$ production is presented. The experimental data complete the measurement of strange particles produced in Al+Al collisions at 1.9$A$ GeV measured with the FOPI detector at SIS/GSI. The $K^*(892)^0$ / $K^0$ yield ratio is found to be $0.0315\pm 0.006 (\mathrm{stat.})\pm 0.012 (\mathrm{syst.})$ and is in good agreement with the UrQMD model prediction. These measurements provide information on in-medium cross section of $K^+$ - $\pi^-$ fusion which is the dominant process on sub-threshold $K^*(892)^0$ production.Comment: 5 pages, 4 figures, accepted for publication in Phys. Rev.

K^+ production in the reaction 58Ni+58Ni^{58}Ni+^{58}Ni at incident energies from 1 to 2 AGeV

Semi-inclusive triple differential multiplicity distributions of positively charged kaons have been measured over a wide range in rapidity and transverse mass for central collisions of $^{58}$Ni with $^{58}$Ni nuclei. The transverse mass ($m_t$) spectra have been studied as a function of rapidity at a beam energy 1.93 AGeV. The $m_t$ distributions of K^+ mesons are well described by a single Boltzmann-type function. The spectral slopes are similar to that of the protons indicating that rescattering plays a significant role in the propagation of the kaon. Multiplicity densities have been obtained as a function of rapidity by extrapolating the Boltzmann-type fits to the measured distributions over the remaining phase space. The total K^+ meson yield has been determined at beam energies of 1.06, 1.45, and 1.93 AGeV, and is presented in comparison to existing data. The low total yield indicates that the K^+ meson can not be explained within a hadro-chemical equilibrium scenario, therefore indicating that the yield does remain sensitive to effects related to its production processes such as the equation of state of nuclear matter and/or modifications to the K^+ dispersion relation.Comment: 24 pages Latex (elsart) 7 PS figures to be submitted to Nucl. Phys