In the past, the role of light as an energy source was largely ignored in research focused on cold acclimation and freezing tolerance in plants. However, cold acclimation is an energy-requiring process. We summarize research illustrating that photoautrophs as diverse as cyanobacteria (Plectonema boryanum), green algae (Chlorella vulgaris, Dunaliella salina, Chlamydomonas raudensis), crop plants (Triticum aestivum L., Secale cereale L., Brassica napus L.), and conifers (Pinus banksiana) L.) tailor the structure and function of the photosynthetic apparatus to changes in temperature and irradiance to maintain cellular energy balance called photostasis. Modulation of either temperature or irradiance results in a similar imbalance in cellular energy that is sensed through changes in chloroplastic excitation pressure. Thus, concepts of photostasis and excitation pressure provide the context through which one can explain the congruence of phenotypic plasticity and photosynthetic performance associated with cold acclimation and photoacclimation. Photosynthetic organisms can sense changes in temperature and irradiance through modulation of the redox state of the photosynthetic electron transport chain, which, in turn, governs phenotype through the regulation of nuclear gene expression and chloroplast biogenesis. We suggest that elucidation of the molecular mechanism(s) by which excitation pressure regulates phenotypic plasticity and photosynthetic performance will be essential in addressing the challenge of maintaining or perhaps enhancing crop productivity under the suboptimal growth conditions predicted to occur as a consequence of climate change.
Sunlight, the ultimate energy source for life on our planet, enters the biosphere as a direct consequence of the evolution of photoautotrophy. Photoautotrophs must balance the light energy absorbed and trapped through extremely fast, temperature-insensitive photochemistry with energy consumed through much slower, temperature-dependent biochemistry and metabolism. The attainment of such a balance in cellular energy flow between chloroplasts, mitochondria and the cytosol is called photostasis. Photoautotrophs sense cellular energy imbalances through modulation of excitation pressure which is a measure of the relative redox state of Q(A), the first stable quinone electron acceptor of photosystem II reaction centers. High excitation pressure constitutes a potential stress condition that can be caused either by exposure to an irradiance that exceeds the capacity of C, N, and S assimilation to utilize the electrons generated from the absorbed energy or by low temperature or any stress that decreases the capacity of the metabolic pathways downstream of photochemistry to utilize photosynthetically generated reductants. The similarities and differences in the phenotypic responses between cyanobacteria, green algae, crop plants, and variegation mutants of Arabidopsis thaliana as a function of cold acclimation and photoacclimation are reconciled in terms of differential responses to excitation pressure and the predisposition of photoautotrophs to maintain photostasis. The various acclimation strategies associated with green algae and cyanobacteria versus winter cereals and A. thaliana are discussed in terms of retrograde regulation and the "grand design of photosynthesis" originally proposed by Arnon (1982).
We hypothesized that chloroplast energy imbalance sensed through alterations in the redox state of the photosynthetic electron transport chain, measured as excitation pressure, governs the extent of variegation in the immutans mutant of Arabidopsis thaliana. To test this hypothesis, we developed a nondestructive imaging technique and used it to quantify the extent of variegation in vivo as a function of growth temperature and irradiance. The extent of variegation was positively correlated (R 2 = 0.750) with an increase in excitation pressure irrespective of whether high light, low temperature, or continuous illumination was used to induce increased excitation pressure. Similar trends were observed with the variegated mutants spotty, var1, and var2. Measurements of greening of etiolated wild-type and immutans cotyledons indicated that the absence of IMMUTANS increased excitation pressure twofold during the first 6 to 12 h of greening, which led to impaired biogenesis of thylakoid membranes. In contrast with IMMUTANS, the expression of its mitochondrial analog, AOX1a, was transiently upregulated in the wild type but permanently upregulated in immutans, indicating that the effects of excitation pressure during greening were also detectable in mitochondria. We conclude that mutations involving components of the photosynthetic electron transport chain, such as those present in immutans, spotty, var1, and var2, predispose Arabidopsis chloroplasts to photooxidation under high excitation pressure, resulting in the variegated phenotype.
Excitation pressure, measured by chlorophyll fluorescence as 1-qP, is an estimate of the redox state of the PQ pool and can be modulated by both light and temperature. We hypothesized that the redox state of the PQ pool regulates the extent of variegation in immutans. To test this hypothesis, immutans was grown at either 25°C or 12°C with increasing light intensities of 50, 150 and 450 μmol photons m−2 s−1 under either a short day photoperiod (8/16 day/night) or under continuous light. An imaging technique was developed to quantify the kinetics of leaf variegation with concomitant measurements of excitation pressure during steady-state photosynthesis. As the growth light intensity increased at 25°C, the extent of variegation increased from no detectable variegation at 50 μmol photons m−2 s−1 to a maximum of 59% variegation at 450 μmol photons m−2 s−1. In contrast, immutans grown at 12°C exhibited significant variegation even when grown at low light under either a short day photoperiod or continuous light conditions. Since the development of white sectors was correlated with excitation pressure, we conclude that the redox state of the PQ pool regulates the extent of variegation.
Understanding the role of the xanthophyll cycle and elucidating the mechanisms of antenna quenching through the non-photochemical dissipation of excess absorbed energy in the photoprotection of the photochemical apparatus continues to be a major focus of photosynthetic research. In addition to antenna quenching, there is evidence for the non-photochemical dissipation of excess energy through the PS II reaction center. Hence, this photoprotective mechanism is called reaction center quenching. One technique to assess reaction center quenching is photosynthetic thermoluminescence. This technique represents a simple but powerful probe of PS II photochemistry that measures the light emitted due to the reversal of PS II charge separation through the thermally-dependent recombination of the negative charges stabilized on Q− A and Q− B on the acceptor side of PS II with the positive charges accumulated in the S2- and S3-states of the oxygen evolving complex. Changes in the temperature maxima for photosynthetic thermoluminescence may reflect changes in redox potentials of recombining species within PS II reaction centers. Exposure of Synechococcussp. PCC 7942, Pinus sylvestrisL., Arabidopsis thaliana, and Chlamydomonas reinhardtii to either lowtemperatures or to high light induces a significant downshift in the temperature maxima for S2Q− B and S3Q− B recombinations relative to S2Q− A and S3Q− A recombinations. These shifts in recombination temperatures are indicative of lower activation energy for the S2Q− B redox pair recombination and a narrowing of the free energy gap betweenQAandQB electron acceptors. This, in turn, is associated with a decrease in the overall thermoluminescence emission. We propose that environmental factors such as high light and low temperature result in an increased population of reduced QA (Q− A), that is, increased excitation pressure, facilitating non-radiative P680+Q− A radical pair recombination within the PS II reaction center. The underlying molecular mechanisms regulating reaction center quenching appear to be species dependent. We conclude that reaction center quenching and antenna quenching are complementary mechanisms that may function to photoprotect PS II to different extents in vivo depending on the species as well as the environmental conditions to which the organism is exposed.
Maximum photosynthetic capacity indicates that the Antarctic psychrophile Chlamydomonas raudensis H. Ettl UWO 241 is photosynthetically adapted to low temperature. Despite this finding, C. raudensis UWO 241 exhibited greater sensitivity to low‐temperature photoinhibition of PSII than the mesophile Chlamydomonas reinhardtii P. A. Dang. However, in contrast with results for C. reinhardtii, the quantum requirement to induce 50% photoinhibition of PSII in C. raudensis UWO 241 (50 μmol photons) was comparable at either 8°C or 29°C. To our knowledge, this is the first report of a photoautotroph whose susceptibility to photoinhibition is temperature independent. In contrast, the capacity of the psychrophile to recover from photoinhibition of PSII was sensitive to temperature and inhibited at 29°C. The maximum rate of recovery from photoinhibition of the psychrophile at 8°C was comparable to the maximum rate of recovery of the mesophile at 29°C. We provide evidence that photoinhibition in C. raudensis UWO 241 is chronic rather than dynamic. The photoinhibition‐induced decrease in the D1 content in C. raudensis recovered within 30 min at 8°C. Both the recovery of the D1 content as well as the initial fast phase of the recovery of Fv/Fm at 8°C were inhibited by lincomycin, a chloroplast protein synthesis inhibitor. We conclude that the susceptibility of C. raudensis UWO 241 to low‐temperature photoinhibition reflects its adaptation to low growth irradiance, whereas the unusually rapid rate of recovery at low temperature exhibited by this psychrophile is due to a novel D1 repair cycle that is adapted to and is maximally operative at low temperature.
IMMUTANS (IM) encodes a thylakoid membrane protein that has been hypothesized to act as a terminal oxidase that couples the reduction of O2 to the oxidation of the plastoquinone (PQ) pool of the photosynthetic electron transport chain. Because IM shares sequence similarity to the stress-induced mitochondrial alternative oxidase (AOX), it has been suggested that the protein encoded by IM acts as a safety valve during the generation of excess photosynthetically generated electrons. We combined in vivo chlorophyll fluorescence quenching analyses with measurements of the redox state of P700 to assess the capacity of IM to compete with photosystem I for intersystem electrons during steady-state photosynthesis in Arabidopsis (Arabidopsis thaliana). Comparisons were made between wild-type plants, im mutant plants, as well as transgenics in which IM protein levels had been overexpressed six (OE-6×) and 16 (OE-16×) times. Immunoblots indicated that IM abundance was the only major variant that we could detect between these genotypes. Overexpression of IM did not result in increased capacity to keep the PQ pool oxidized compared to either the wild type or im grown under control conditions (25°C and photosynthetic photon flux density of 150 μmol photons m−2 s−1). Similar results were observed either after 3-d cold stress at 5°C or after full-leaf expansion at 5°C and photosynthetic photon flux density of 150 μmol photons m−2 s−1. Furthermore, IM abundance did not enhance protection of either photosystem II or photosystem I from photoinhibition at either 25°C or 5°C. Our in vivo data indicate that modulation of IM expression and polypeptide accumulation does not alter the flux of intersystem electrons to P700+ during steady-state photosynthesis and does not provide any significant photoprotection. In contrast to AOX1a, meta-analyses of published Arabidopsis microarray data indicated that IM expression exhibited minimal modulation in response to myriad abiotic stresses, which is consistent with our functional data. However, IM exhibited significant modulation in response to development in concert with changes in AOX1a expression. Thus, neither our functional analyses of the IM knockout and overexpression lines nor meta-analyses of gene expression support the model that IM acts as a safety valve to regulate the redox state of the PQ pool during stress and acclimation. Rather, IM appears to be strongly regulated by developmental stage of Arabidopsis.
To study the transcriptional regulation of Aox1, the major alternative oxidase (AOX) gene, we fused 1072 bp of its promoter to the promoterless arylsulfatase reporter gene. We find that the reporter is strongly activated when cells are shifted from a medium containing ammonium to one containing nitrate but is unresponsive to treatments known to induce Aox1 in higher plants – H2O2, antimycin A and cold stress. However, induction of Aox1 by all these factors was found when changes in gene expression were monitored using RNA blot analysis. Our data suggest that transcriptional upregulation of Aox1 by H2O2, antimycin A and cold‐stress requires one or more enhancers which are not found in the proximal promoter region of the gene. Interestingly, while nitrate, H2O2 and cold stress result in increased AOX abundance and increased alternative pathway respiration, these changes are not seen when cells are treated with antimycin A. We find that H2O2, antimycin A and cold‐stress result in an increase in intracellular reactive oxygen species (ROS) but that a significant change in ROS does not occur when cells are shifted into a medium‐containing nitrate. Overall, our data are consistent with there being two distinct pathways regulating Aox1 transcription and AOX abundance in Chlamydomonas reinhardtii: one in response to oxidative stress and a second due to metabolic changes brought about by a shift in nitrogen source from ammonium to nitrate.
Recovery from photoinhibition in the low-light adapted psychrophilic green alga, Chlamydomonas raudensis (UWO 241). : 13th International Congress of Photosynthesis
Recovery from photoinhibition in the low light-adapted psychrophilic green alga Chlamydomonas raudensis (UWO 241).
BACKGROUND:Dural cavernous angiomas are uncommon benign vascular malformations which may present intraoperative difficulties in hemostasis when the diagnosis is not suspected preoperatively. Preoperative diagnosis can be difficult when angiomas show atypical features and share imaging characteristics with other entities.METHODS:A patient presented with a radiographically aggressive lesion, subsequently identified as a dural cavernous angioma. The lesion is reviewed and its clinical, radiographic, and pathological features are compared with other vascular malformations.CASE REPORT:A 40-year-old man presented with new onset seizures and an enhancing lesion infiltrating the floor of the right middle cranial fossa. Due to its aggressive radiographic appearance, initial considerations included chondrosarcoma, meningioma or metastasis. Pathological examination, however, revealed the lesion to be a cavernous angioma of dura.CONCLUSION:This uncommon lesion may present a diagnostic challenge with significant intraoperative implications. T2 sequence hyperintensity in a relevant lesion should raise suspicion of an hemangioma. It is important to be aware of this entity and its potential to mimic other entities on radiographic grounds.
We present a unique case of a patient with a symptomatic carotid cavernous fistula treated successfully with balloon embolization. Her subsequent death from other disease processes allowed direct visualization of the balloon occlusion in situ at postmortem examination.
We describe a technique used to treat two patients with large, wide-necked aneurysms during the past 2 years. In the initial attempts at embolization, evidence of coil instability within the aneurysm or significant impingement of coil loops on the parent artery was observed. Advancement of a second microcatheter into the aneurysm allowed two coils to be braced across the aneurysmal neck before the detachment of either coil. This technique permitted successful coil treatment in both patients.
A 42 yr old male presented with left facial weakness. MRI showed lesions affecting the distal seventh nerve and third division of the trigeminal nerve. The seventh nerve was biopsied and showed a malignant epithelioid schwannoma. The patient underwent extensive resection followed by irradiation. This is one of very few examples of intracranial malignant peripheral nerve sheath tumors and the first reported example of an intracranial malignant epithelioid schwannoma. The literature is reviewed and completeness of resection appears to be the most pertinent prognostic factor.
ABSTRACT: Background: Intracranial dural arteriovenous fistula with pial venous drainage may present with hemorrhage or focal neurologic deficit and may be difficult to treat. We wish to summarize the therapeutic approaches to these potentially dangerous lesions and to demonstrate how endovascular and neurosurgical therapies may have complimentary roles in their management. Methods: The clinical and radiological records of all patients who presented to our institution with intracranial dural arteriovenous fistula over the last 5 years were reviewed. In those cases demonstrating pial venous drainage, details of presentation, imaging features, endovascular and surgical therapy and outcome were analyzed. Results: We identified 13 patients with these lesions, 7 of whom presented with intracranial hemorrhage. Six patients were treated with embolization alone. Angiographic cure was achieved in 4. There was one complication in this group, a subarachnoid hemorrhage following glue injection. Four patients were treated with embolization followed by surgical occlusion of the pial venous drainage. Angiographic cure was achieved in all 4. There was one complication in this group, a facial nerve palsy following glue injection. Three patients were treated by surgery alone, with no complications and complete cure in all. Conclusion: Endovascular therapy of intracranial dural arteriovenous fistula may be curative but is often complex and carries definite risks. Neurosurgical ligation of pial draining veins, with pre-operative embolization when safe, may be a relatively more controlled method to achieve complete cure.