Imipramine affects circadian leaf movements in Oxalis regnellii

The sensitivity of the circadian leaf movement of Oxalis regnellii Mig. to imipramine (a tricyclic dibenzazepine) was investigated. Imipramine, like Li⁺, is used as a therapeutic agent against depressive disorders in man. The therapeutic effects of the two substances might be mediated via effects on basic circadian rhythms and the cellular level. It was indeed possible to influence the circadian movement of Oxalis by imipramine; pulses (10⁻³ M , 4h) phase shifted the rhythms and caused advances. A phase response curve is presented. No period change of the movements was caused by permanent presence of imipramine (5 - 10⁻⁵ or 10⁻⁵ M ). The nature of the imipramine-induced phase shift is discussed and compared with Li⁻¹ effects on the same circadian system.     

A method to record circadian plant movements, with application to Oxalis leaf rhythms

A transducer was developed to record the circadian movement of the individual leaflets in Oxalis regnellii Mig. The method can easily be adapted to measure other kinds of plant movements as well. It is based on the detection of the shadow each leaflet casts on the small side of a specially formed Perspex plate. The light is guided through the Perspex and collected by a phototransistor, which provides an electrical signal that is proportional to the light intensity falling onto it. The output signal can be made a linear function of the leaf angle. This equipment was used in experiments to study the coupling between the 3 leaflets in Oxalis . Pulses of 4 h of red light were given to one of the leaflets, the two others were shielded from the light. A phase response curve was determined for each leaflet, but there was no significant difference in the phase response between the 3 leaflets. Experiments were also made in which the 3 leaflets were separated physically by cuts along the petiole between the pulvini. In this case ultradian oscillations were observed.     

Effect of abscisic acid on the circadian leaf movements of Oxalis regnellii

The effect of abscisic acid on the circadian leaf movements of Oxalis regnellii Mig. was investigated. Leaf stalks were cut and put into a nutrient solution, and the leaf movements automatically recorded. Continuous application of ABA (10^-5 or 10^-4 M ) to the solution did not alter the circadian period. However, the treatment delayed the rhythm and the amplitude was somewhat decreased. ABA pulses (4 h, 10^-4 M ) caused permanent phase shifts of the rhythm with delays and advances. Transient wilting of the stalks plus leaves did not produce permanent phase shifts of the rhythm. A wilting period of 4 h gave a (transient) 2 h advance as judged from the first maximum after the wilting treatment. This advance was independent of the time of application of the wilting. Wilting treatment for 8 h caused advances, the magnitude of which varied according to the timing of the wilting treatment. It is concluded that there was no connection between the (permanent) ABA effects and the (transient) wilting effects on the phase of the circadian rhythm. A direct action of ABA on the circadian oscillator is possible and the molecular mechanism might be via effects on ion permeability of the pulvini cells.     

Basic properties of the circadian leaf movements of Oxalis regnellii, and period change due to lithium ions

The circadian leaf movement of Oxalis regnellii Mig, has been investigated. The three leaflets of a stalk were normally synchronized, and under the experimental conditions chosen they showed a period of 26.2 ± 0.1 h. Cutting off one or two leaflets led to a successive decrease of the period length (25.7 ± 0.1 and 25.1 ± 0.3 h resp.). It was possible to phase shift the leaf movements by mechanical means (advance of 1.6 ± 0.3 h).
Lithium ions, added permanently to the transpiration stream, increased the period length of the leaf movements by more than one hour (with 10 m M Li⁺). A 24 h pulse of 20 m M LiCl caused a permanent 2–3 h phase delay of the circadian rhythm. Four-h pulses, on the other hand, provoked only transient phase delays, the magnitude being dependent on the phase of application. Lithium concentrations were determined for different regions of leaves and pulvini. It was shown that leaf segments had considerably lower concentrations than pulvini. No significant difference in the lithium concentration was observed between the upper and lower part of pulvini.
In the light leaf position was strongly correlated with water uptake and the consequences for applications of substances to the circadian system via the transpiration stream is discussed. A simple model of the oscillatory system and reactions connected to it is discussed.     

Effects of alkali ions on the circadian leaf movements of Oxalis regnellii

The influence of alkali ions on the circadian leaf movements of Oxalis regnellii Mig. was investigated. Ions were given to the oscillating system via the transpiration stream of cut stalks in nutrient medium. Chloride solutions of Rb⁺, Cs⁺, Na⁺ and K⁺ were tested and the results compared to previously published LiCl-results. The period of the circadian leaf movements was unaffected by a continual addition of Na⁺ or K⁺ to the nutrient medium (at least up to 40 mM). Rb⁺, in the concentration of 2.5 or 5 mM, caused a shortening of the period when applied continuously. Rb⁺ concentrations up to 60 mM were tested. Cs⁺ ions caused only lengthenings of the circadian period. Cs⁺ concentrations up to 40 mM were tested. Cs⁺ resembled Li⁺ in producing period lengthenings, but was not as effective as Li⁺ when compared on a concentration basis. Toxicity of the effective ions was in the following order: Li⁺Cs⁺Rb⁺, Rb⁺ pulses (50 mM, 4 h) phase-shifted the rhythm and caused advances. A phase response curve was determined and the maximum steady state advances were of the order of 1 h. The dual effect of the Rb⁺ ions is discussed and is assumed to be due to two counteracting processes, exemplified by Rb⁺-sensitive ATPase-controlled pumping processes and protein synthesis. For comparison, the effects of Rb⁺ and Li⁺ in human depressive disorders is also discussed in relation to their influence on circadian systems. It is emphasized that Rb⁺ and K⁺ behave differently and are not interchangeable in their action on circadian systems.     

酢浆草属5种植物的微形态特征观察

利用光镜和扫描电镜观察了酢浆草属(Oxalis L.)5种植物的叶、花、花粉和部分种子的微形态结构。5种植物叶片上均具有平列型气孔器,外围3~4个不规则表皮细胞;酢浆草(O.corniculata L.)、铜锤草(O.corymbosa DC.)及白花酢浆草(O.acetosella L.)叶片两面均有气孔器,下表皮犹密;紫叶酢浆草(O.triangularis A.St.-Hil.)和山酢浆草(O.griffithii Edgew.et Hook.f.)的气孔器只分布于叶片下表皮。不同种间及长、短雄蕊上的花粉粒大小各异,花粉多呈近球形(酢浆草、白花酢浆草)、长球形(紫叶酢浆草、山酢浆草)或超长球形(铜锤草),极面观为三裂圆形,赤道面大多具3沟,少数4沟,表面具有不规则的穴状或粗网状纹饰,网眼内无或有乳突状突起(铜锤草、紫叶酢浆草)。酢浆草种子较小,表皮纹饰为比较规则的不等边六角形网眼,网眼中间具棱柱状突起,上有钩状附属物;山酢浆草种子较大,表皮粗糙程度低于酢浆草。     

Anatomical and physiological characteristics of the petiole of Abutilon theophrasti in relation to circadian leaf movements

Leaf movements in Abutilon theophrasti Medic, were monitored manually and by a continuous electronic recording device. Plants entrained to a daily regime of a 15 h light span followed by 9 h of darkness showed rhythmic movements that persisted under conditions of continuous illumination and constant temperature with a circadian period. The rhythmic change in orientation of the leaf from a near horizontal (day) to a near vertical (night) position was attributed to movement of the blade and not the petiole. The end of the petiole next to the blade functions as a joint or pulvinus. Anatomical confirmation of the existence of a pulvinus in the Abutilon leaf was provided by light microscopy. Vascular tissue in this region forms a solid cylinder with no pith, and the cortex is parenchymatous. In the main part of the petiole, the vascular tissue is arranged in four to six bundles, a pith is present, and the cortex contains a sub-epidermal ring of collenchyma. Both the functional and anatomical evidence indicate the presence of a pulvinus that functions in circadian leaf movements of Abutilon.     

Self‐excitation of leaf movements in Oxalis regnellii under different light qualities

Abstract

Self‐excitation of leaf movements in Oxalis regnellii was studied under the influence of different constant conditions as darkness or irradiation of red or far‐red. Since in Oxalis two oscillating systems have to be regarded to, attention was focussed upon the development in time of self‐excitation of both systems. Similar to previous findings under dim white light the self‐excitation of the spreading system preceeds the foldings system in time in all cases.

A behavioral difference as response to red and far‐red could be found in the rate of self‐excitation. While under the influence of continuous red light maximal amplitudes are reached within three free‐running periods, under far‐red this effect needs only one period. In connection to this result, the deceleration of leaf closure caused by far‐red (well‐known from experiments in legumes, and now also observed in Oxalis), can be shown to reflect the interference of two antagonistic effects. Since self‐excitation starts rapidly under far‐red and generally begins with the excitation of the spreading movement, nastic closure can be compensated by the first free‐running spike of the spreading system. In addition, a computer program is presented to simulate amplitude‐dips and phase‐shifts caused by antagonistic effects of two locally separated systems. In this program each oscillator activity is implied to perform a steep spike with following slow decline (collision function).     

Changes in leaf optical properties associated with light-dependent chloroplast movements

We surveyed 24 plant species to examine how leaf anatomy influenced chloroplast movement and how the optical properties of leaves change with chloroplast position. All species examined exhibited light-dependent chloroplast movements but the associated changes in leaf absorptance varied considerably in magnitude. Chloroplast movement-dependent changes in leaf absorptance were greatest in shade species, in which absorptance changes of >10% were observed between high- and low-light treatments. Using the Kubelka-Munk theory, we found that changes in the absorption (k) and chlorophyll a absorption efficiency (k*) associated with chloroplast movement correlated with cell diameter, such that the narrower, more columnar cells found in sun leaves restricted the ability of chloroplasts to move. The broader, more spherical cells of shade leaves allowed greater chloroplast rearrangements and in low-light conditions allowed efficient light capture. Across the species tested, light-dependent chloroplast movements modulated leaf optical properties and light absorption efficiency by manipulating the package (sieve or flattening) effect but not the detour (path lengthening) effect.     

Effects of vanadate, N2 and light on the membrane potential of motor cells and the lateral leaflet movements of Desmodium motorium

The lateral leaflets of Desmodium motorium (Houtt.) Merr. exhibit ultradian up- and down movements, which are paralleled by oscillations of the membrane potential of motor cells in the pulvinus. By different treatments we have tested the hypothesis that both that both oscillation-types are causally related. The reactions of the leaflet movement and the membrane potential were evaluated by the following approaches. (1) Application of vanadate. an inhibitor of the proton pump in the plasmalemma. and N₂ suppressed leaflet movements and finally arrested the leaflet in the lower position. Before the oscillations damped out, a strong lengthening in period was found. This indicates that the pump is part of the ultradian clock. A period lenthening and a final suppression of the rhythm by vanadate was also seen in the extracellular electric potential of the pulvinus. Intracellular recordings in situ showed that vanadate application depolarized the motor cells. (2) Light of high fluence rates diminished the amplitude of the oscillations of the membrane potential of single motor cells and shortened the period. The same effects were observed when monitoring the lateral leaflet movement. The leaflet always moved towards the direction of the light. whether it was applied from the abaxial or from the adaxial part of the pulvinus. (3) When light was applied to the pulvinus of lateral leaflets. which had spontancously stopped moving in an upper position. oscillations were induced transiently. This effect was also found for the membrane potential of motor cells in the pulvinus. - Our results thus provide further evidence that the membrane potential controls the volume state of the motor cells in the pulvinus of lateral leaflets of Desmodium motorium .     

Circadian Modulation of Cold-Induced Thermogenesis in the Golden Hamster

The relationship between the circadian and homeostatic control of body temperature was studied in golden hamsters maintained under a 14:10 LD cycle. Telemetric records of body temperature showed that body temperature oscillates daily with a low phase during the light section of the LD cycle and a high phase during the dark section. The low phase of the temperature rhythm was found to start two hours after lights on and to last about 8 hours. The high phase was found to start immediately after lights off and to last about 8 hours also. Metabolic heat production was measured by indirect calorimetry during the high phase and the low phase of the body temperature rhythm. Heat production in a thermoneutral environment was higher during the high phase of the body temperature rhythm than during the low phase, but cold-induced thermogenesis was greater during the low phase than during the high phase. This finding suggests that the autonomic thermoregulatory system is more responsive to cold stress during the low phase than during the high phase. Consequently, the daily oscillation of body temperature cannot be explained by an elevation of the thermoregulatory set point during the high phase of the rhythm. The homeostatic and circadian control of body temperature seem to be exerted separately from each other.     

Aquaporins and plant leaf movements

BaCKGROUND: Plant leaf movements can be mediated by specialized motor organs, the pulvini, or can be epinastic (i.e. based on different growth velocities of the adaxial and abaxial halves of the leaf). Both processes are associated with diurnally regulated increases in rates of membrane water transport, which in many cases has been shown to be facilitated by aquaporins. Rhythmic leaf movements are known from many plant species, but few papers deal with the involvement of aquaporins in such movements. SCOPE: Many details of the architecture and function of pulvini were worked out by Ruth Satter and co-workers using Samanea saman as a model organism. More recently a contribution of aquaporins to pulvinar movement in Samanea was demonstrated. Another model plant to study pulvinus-mediated leaf movements is Mimosa pudica. The contribution of both plasma membrane- and tonoplast-localized aquaporins to the seismonastic leaf movements in Mimosa was analysed. In tobacco, as an example of epinastic leaf movement, it was shown that a PIP1 aquaporin family member is an important component of the leaf movement mechanism.     

Endogenous patterns of photomechanical movements in teleosts and their relation to activity rhythms

Summary The retinal rods, cones and epithelial pigment of most lower vertebrates display rhythmic photomechanical (retinomotor) migrations in response to changes in ambient lighting conditions. This study examines the extent of these migrations in the absence of the daily changes in illumination (constant darkness and constant light) in three species of teleosts. Salmo trutta, a crepuscularly active fish, showed two peaks of light adaptation occurring around dawn and dusk when kept in constant darkness. Tinca tinca, a nocturnal species, also showed an endogenous rhythm during extended periods of darkness, but, unlike Salmo trutta, it was light-adapted throughout what would normally have been day. At the maximal extent of migration under conditions of continual darkness, the pigment migrated 59% as much as it did during a normal light/dark cycle. Nannacara anomala, a tropical diurnally active species, showed a similar but more pronounced rhythm than Tinea tinea for all 3 days of experimental darkness, behaving essentially identically to fish exposed to a light/dark cycle. Nannacara anomala also displayed a weak rhythm when kept in constant light.It is concluded from these and previous results that the pattern of endogenous photomechanical movement depends both on the activity pattern of a species and on the constancy of the lighting conditions to which it has been exposed during its lifetime.     

Epidermal focussing and effects on light utilization in Oxalis acetosella

lmage analysis was used to quantify the lateral heterogeneity of the radiation field within the palisade of Oxialis acetosella L. leaves. Oxalis acetosella epidermal cells focus light up to four times incident irradiance, resulting in regions of high and low internal fluenec rate within the palisade. Chlorophyll fluorescence from leaves irradiated with directional light was found to originate primarily from palisade cell chloroplasts located within focal zones. When the internal radiation field was made more homogeneous by using diffuse light or by coating the leaf with a layer of mineral oil to eliminate epidermal focussing, the characteristics of the chlorophyll fluorescence signal were altered: non-photochemical quenching (qN) increased, while relaxation of qN was slowed. This indicates that upper palisade chloroplasts may fine-tune their light utilization to the intra leaf light microenvironment.     

Les rythmes endogènes des mouvements foliaires chez Mimosa pudica L.

Abstract

Endogenous Rhythms in Mimosa pudica L. Leaf Movements.

The rhythmic movements performed by the leaves of the “Sensitive plant”, Mimosa pudica L., observed by time lapse photography, result of periodical turgor variations taking place in the parenchymatous cells of specialized motor organs. These turgor variations are associated with membrane permeability changes and ionic movements. These leaf movements allow to specify the temporal organization of this plant. Statistical analysis of observed periodicities in leaf movement shows that, in alternating conditions of light and dark (L/D:14/10) three distinct rhythms exist: a circadian rhythm synchronized by the photoperiodic cycle (τ = 24 hrs), and two ultradian rhythms with mean period values 3.8 hrs and 0.5 hrs respectively. In constant conditions from germination (L/L), the leaf behavior is strongly modified, but the three period values are found again (mean period values of 25.1 hrs, 3.5 hrs and 0.6 hrs respectively). The occurence of many rhythms with various periods taking place in the same organ is discussed in reference to observations effected on other biological subjects. Then, it appears that the period value within 2 and 4 hrs may be considered as a characteristic one in plants.     

The effect of lunisolar tidal acceleration on stem elongation growth,nutations and leaf movements in peppermint (Mentha × piperita L.)

• Orbital movement of the Moon generates a system of gravitational fields that periodically alter the gravitational force on Earth. This lunar tidal acceleration (Etide) is known to act as an external environmental factor affecting many growth and developmental phenomena in plants. Our study focused on the lunar tidal influence on stem elongation growth, nutations and leaf movements of peppermint.
• Plants were continuously recorded with time‐lapse photography under constant illumination as well in constant illumination following 5 days of alternating dark–light cycles. Time courses of shoot movements were correlated with contemporaneous time courses of the Etide estimates. Optical microscopy and SEM were used in anatomical studies.
• All plant shoot movements were synchronised with changes in the lunisolar acceleration. Using a periodogram, wavelet analysis and local correlation index, a convergence was found between the rhythms of lunisolar acceleration and the rhythms of shoot growth. Also observed were cyclical changes in the direction of rotation of stem apices when gravitational dynamics were at their greatest. After contrasting dark–light cycle experiments, nutational rhythms converged to an identical phase relationship with the Etide and almost immediately their renewed movements commenced. Amplitudes of leaf movements decreased during leaf growth up to the stage when the leaf was fully developed; the periodicity of leaf movements correlated with the Etide rhythms.
• For the fist time, it was documented that lunisolar acceleration is an independent rhythmic environmental signal capable of influencing the dynamics of plant stem elongation. This phenomenon is synchronised with the known effects of Etide on nutations and leaf movements.

     

Shade avoidance and the regulation of leaf inclination in Arabidopsis

As a rosette plant, Arabidopsis thaliana forms leaves near to the ground, which causes the plant to be vulnerable to shading by neighbours. One mechanism to avoid such shading is the regulation of leaf inclination, such that leaves can be raised to more vertical orientations to prevent neighbouring leaves from overtopping them. Throughout Arabidopsis rosette development, rosette leaves move to more vertical orientations when shaded by neighbouring leaves, exposed to low light levels or placed in the dark. After dark-induced reorientation of leaves, returning them to white light causes the leaves to reorient to more horizontal inclinations. These light-dependent leaf movements are more robust than, and distinct from, the diurnal movements of rosette leaves. However, the movements are gated by the circadian clock. The light-dependent leaf orientation response is mediated primarily through phytochromes A, B and E, with the orientation varying with the ratio of red light to far-red light, consistent with other shade-avoidance responses. However, even plants lacking these phytochromes were able to alter leaf inclination in response to white light, suggesting a role for other photoreceptors. In particular, we found significant changes in leaf inclination for plants exposed to green light. This green light response may be caused, in part, by light-dependent regulation of abscisic acid (ABA) biosynthesis.     

Changes in reciprocal herkogamy during the tristyly-distyly transition in Oxalis alpina increase efficiency in pollen transfer

Although the spatial separation of sexual organs within a flower (herkogamy) has been interpreted as a mechanism that promotes efficient pollen transfer, there have been few attempts to relate variation in herkogamy to probabilities of pollen flow. Here, we used a heterostylous species with variation in reciprocal herkogamy to test this hypothesis. We measured legitimate and illegitimate pollen flow with fluorescent dyes in four selected populations of Oxalis alpina corresponding to the extremes of a previously reported evolutionary gradient from tristyly to distyly. After the breakdown of tristyly, the observed increment in reciprocal herkogamy between the long and short morphs was associated with a 30% increase in the proportion of dye received from compatible illegitimate pollinations. In all populations, the most likely effective pollen vectors were two Heterosarus bee species. Our results support the adaptive value of reciprocal herkogamy in promoting efficient pollen transfer in heterostylous species.     

12-hour phase-shift of mice kidney rhodanese (thiosulfate sulfurtransferase) activity in the first two months of life

This study aimed to investigate and compare variation of renal rhodanese activity at 2^nd, 4^th and 8^th weeks of post-natal development (PND) in mice. The enzyme activity increased with age and was higher in females compared to males in all studied groups. Cosinor analysis revealed significant circadian rhythms (with period τ = 24 h) of enzyme activity in both genders with peak time shift during the PND. At the 2^nd week of PND (pre-weaning time), the circadian rhythm peaked at the beginning of light span, more precisely ≅1 HALO (Hours After Light Onset). A week after weaning (4^th week of PND), the peak time was located at the second half of photophase (≅9 HALO) in both genders. Four to six weeks later, about the 8^th week of PND, the circadian peak time was then recorded at ≅13 HALO. These findings suggest that rhodanese level and rhythm stabilization were age-dependent. Moreover, gender-related differences may stimulate discussions on the relationship between renal rhodanese and cyanide sensitivity.