Studies in Stomatal Behaviour: V. THE ROLE OF CARBON DIOXIDE IN THE LIGHT RESPONSE OF STOMATA

It was found that stomata on illuminated leaves, both of Pelargoniumand wheat, opened much wider where the leaf surface was enclosedin a small volume of air, as in a normal porometer cup, thanelsewhere. This was shown for both species by the infiltrationmethod, and for Pelargonium by Lloyd's method and direct microscopicalobservation also. The effect was shown not to be due to pressure of the porometercup or glass plate on the leaf, or to temperature differences,nor directly to the lack of movement or high humidity of theenclosed air. A considerable body of data was collected which appeared tosupport the hypothesis that the wide opening was due to accumulationof some volatile substance produced by the leaf, but all theresults were also consistent with the view that it was causedby reduction in the carbon dioxide content of the enclosed airbelow the normal 0·03 per cent. owing to photosynthesis.Further crucial experiments with both the porometer and infiltrationmethods left virtually no doubt that the latter hypothesis wascorrect. This extreme sensitivity of stomata to carbon dioxide concentrationwithin the range 0·03 per cent. to zero is discussedin relation to their operation in nature, and a possible biologicaladvantage is suggested. The bearing of the effect upon porometer investigations is alsodiscussed and it is concluded that for all quantitative or semi-quantitativeexperimentation it is essential to use a cup detached betweenreadings, or at least swept with air such as surrounds the restof the leaf, and to have the upper leaf surface above the cuparea freely exposed or similarly swept. For qualitative investigationof the light response of stomata the traditional form of cupmay be used. The importance is stressed of allowing porometer readings toreach equilibrium under one set of conditions before changingto another, when investigating the ‘closing’ or‘opening’ effects of external factors. Several subsidiary effects, observed in the course of the investigation,are discussed; in particular an effect of humidity upon therate of response to other factors.     

Studies in Stomatal Behaviour: VI. AN INVESTIGATION OF THE LIGHT RESPONSES OF WHEAT STOMATA WITH THE ATTEMPTED ELIMINATION OF CONTROL BY THE MESOPHYLL

As in the previous investigation (Part I), stomatal responsesin wheat were followed by means of porometer readings; betweenthese, air was forced through the stomata to sweep out the substomatalcavities. Instead of forcing the whole of a relatively slowair-flow in through the stomata, however, a fast flow was sweptacross the leaf surface and a portion of this passed into theintercellular space system. This helped to maintain the desiredcarbon dioxide content close to the stomatal openings and thestomata could thus be treated with different carbon dioxideconcentrations. A factorial experiment was carried out in which6 carbon dioxide concentrations from zero to 0084 per cent,were combined with 3 light intensities from 90 to 800 f.c, the18 treatments being replicated 6 times in three ‘66 latinsquares’. A supplementary experiment in which the same6 carbon dioxide treatments were given in darkness was carriedout in duplicate. The results confirmed previous findings inthat the stomata showed no appreciable response to variationof external carbon dioxide supply between zero and 001 percent, concentration and that a large and highly significantlight response was found at each of these two carbon dioxidelevels. This is held to indicate the existence of a light effect(direct or indirect) not operating by reduction of the internalcarbon dioxide content of the guard cells. On the assumptionthat the rates of air flow through the stomata were adequate,the combined results of the two experiments are interpretedmainly in terms of two direct effects of light upon the stomata,one depending on photosynthetic reduction of carbon dioxidein the guard cells and the other, mentioned above, not so operating.To these must be added the indirect effect responsible for the‘transmission’ described in Part 1, which mightpossibly account for the whole of the second direct effect postulated.Finally, if the rates of flow were in fact grossly inadequate,the trends attributed to the first direct effect mentioned abovemight have been due to indirect control by the mesophyll viathe intercellular space atmosphere. Such control must accountfor part of the light responses which occur under natural conditions.The natural light response is thus made up of at least two andmore probably four components—two direct and two indirect,in each case one operating by the internal carbon dioxide andthe other not. A highly significant diurnal rhythm of stomatalmovement under constant illumination and temperature is shownto occur in wheat. The measure of stomatal resistance used (logresistance per 10,000 stomata) is discussed, and it is concludedthat although it is empirical a much more satisfactory measureis not at present available.     

The Influence of Water Strain on the Minimum Intercellular Space Carbon dioxide Concentration {lambda} and Stomatal Movement in Wheat Leaves

Eight detached wheat leaves were each subjected to varying degreesof controlled water strain, in a specially constructed leafchamber (which is described in detail) connected to a closedsystem incorporating an infra-red gas analyser. The water strainwas varied by applying successively to their leaf sheaths twoconcentrations of mannitol solution (0·2 M. and 0·4M.) each preceded and followed by water. Conditions were maintainedconstant for light (900 f.c.), leaf temperature (25° C.)and humidity (50 per cent. RH) of the circulating air as itentered the leaf chamber. Steady-state values (gamma) of thecarbon dioxide concentration in the closed system were recordedboth for the water and mannitol treatments and the data analysedstatistically. The two mannitol treatments were found to increasegamma from 80 p.p.m. with water to 94 and 116 p.p.m. respectively,all differences being highly significant. Concomitant measurementswere made of stomata1 resistance with a porometer and of transpirationby means of wet and dry thermocouples; the stomata showed markedclosure in response to mannitol treatment (complete closurewith the higher concentration) and partial reopening with theleaf sheath in water once more, transpiration also falling andrising inversely with water strain. It is concluded that part of the closing response of wheat stomatato water strain under natural conditions must be operated byincrease in the intercellular-space carbon dioxide concentration,but a comparison with previous experimental results suggeststhat this can only be a small part unless sensitivity to carbondioxide is greatly increased by water loss. The initial response to mannitol treatment was a preliminaryopening of the stomata and a fall of about 10 per cent. in carbondioxide concentration. These results imply that under naturalconditions the preliminary opening of wheat stomata with wiltingmay be partly passive and partly active with a carbon dioxide-operatedmechanism. A subsidiary experiment with water strain varied by removingand restoring the water supply to the leaf sheath gave increasesand decreases in carbon dioxide similar to those in the mainexperiment; the results of the latter need not, therefore, beattributed to metabolism of the mannitol. It is suggested that values of gamma give a good measure ofthe efficiency of the mesophyll for net carbon dioxide absorption,with the stomatal factor eliminated, under the given conditionsand with carbon dioxide ‘limiting’ in the senseused by Maskell (1928).     

Studies in Stomatal Behaviour: VIII. STOMATAL RESPONSES TO TEMPERATURE AND CARBON DIOXIDE CONCENTRATION IN ALLIUM CEPA L., AND THEIR RELEVANCE TO MID-DAY CLOSURE

It has been reported in the literature that onion leaf stomatatend to close at noon (Loftfield, 1921) and that the minimumcarbon dioxide concentration () in onion leaf tissue rises from0.012 per cent to 0.024 per cent. When the leaf temperatureincreases from 30°C. to 35°C (light intensity 900.f.c.)(Heath and Orchard, 1957). Experiments were therefore carried out of test the hypothesisthat raising the left temperature above about 30°C. causesstomatal closing movements in onion leaves (e.g. mid-day closure)by increasing the carbon dioxide concentration in the leaf tissue.This hypothesis has been supported by the results of these experiments.Another and diametrically opposed effect of temperature on stomatalmovements in onion leaves has, however, been discovered, Namelythat, provided the final width of stomatal pores.     

THE LABELLING OF THE GANGLIOSIDIC FRACTION FROM BRAINS OF RATS EXPOSED TO DIFFERENT LEVELS OF STIMULATION AFTER INJECTION OF [6-3H]-GLUCOSAMINE

[6-³H] Glucosamine was injected into rats and then they were submitted for 1 h to light (about 1100 lux) and sound, while their controls were kept in darkness. Two series of experiments were done: in one, the animals received the injection intracerebrally; in the other, the injection was intraperitoneal. In both series the ratios of specific radioactivities (c.p.m./μmol of NANA) in mitochondria1 gangliosidic fractions to those in synaptosomal gangliosidic fractions were higher for rats exposed to illumination and sound ?+ 74 per cent, P < 0·01 and + 49 per cent, P < 0·001 (in respectively, experiments with intracerebral and intraperitoneal injections) than for animals kept in dark. For animals iqjected intraperitoneally the specific radioactivity of the mitochondria1 gangliosidic fraction from rats exposed to light was higher (+ 11 per cent, P < 0.05) than that obtained from animals kept in dark. For synaptosomes the specific radioactivity of the gangliosidic fraction obtained from animals exposed to light was lower (?15 per cent, P < 0.01) than that obtained from animals kept in dark. In animals kept under the experimental conditions already described for more than 1 h the differences tended to disappear. No differences were observed between the two groups of rats in the amounts of radioactivity found in the brain.     

THE EFFECT OF CARBON DIOXIDE TENSION ON THE METABOLISM OF CEREBRAL CORTEX AND MEDULLA OBLONGATA

Manometric measurements were made of oxygen uptake (Q _OO2) and aerobic lactic acid output (Q_G) by slices of cerebral cortex and medulla oblongata of the cat in the presence of mixtures of 1, 5, and 20 volumes per cent of carbon dioxide in oxygen. The concentrations of NaHCO₃ and NaCl in the medium were varied to maintain constant pH and sodium ion concentrations. The calcium ion concentration was 0.0002 M. At pH 7.5 under these conditions, an increase in carbon dioxide from 1 per cent to 5 per cent doubled the Q_G of both tissues but did not alter Q _OO2; an increase from 5 per cent to 20 per cent carbon dioxide had no further effect on Q_G in either tissue or Q _OO2 of cortex, but did depress the Q _OO2 of medulla. At pH 8.1, an increase in carbon dioxide from 1 per cent to 5 per cent raised the Q _OO2 and Q_G of cortex by about 60 per cent. Measurements at low oxygen tension carried out previously in phosphate medium were repeated in bicarbonate medium to obtain data for the combined output of lactic acid and carbon dioxide (Q_A). When the oxygen in the gas phase was decreased from 95 to 3 volumes per cent, the lactic acid output as measured colorimetrically increased by 114 mg./gm. in cortex and by 8 mg./gm. in medulla; Q_A increased from 12.3 to 13.5 in cortex and decreased from 5.1 to 3.8 in medulla.     

An Examination of the Factors affecting Variability in the Growth of the Mesocotyl and Coleoptile of Etiolated Avena Seedlings

Experiments were carried out on the effect of rate of air-flowand of humidity on the growth of Avena seedlings. The experimentalmethods used are described in some detail. The effect of air-flow on mesocoryl growth was considerablewhen unsaturated air was used, but with saturated air growthwas independent of air-flow. The effect must therefore be dueto variation in transpiration rate. A 10 per cent, variationin relative humidity, at a constant rate of flow, caused a 30per cent, variation in mesocotyl length. The coleoptiles showedless pronounced effects, but even in saturated air variabilitywas not eliminated. Under the conditions used a difference of 2·3 mm. inmesocotyl length (per 50 plants), i.e. 8 per cent., was significantat P = 0·05; for the coleoptiles a difference of 1 mm.(8 per cent.) was significant at this level.     

The Composition of the Gas contained in the Vesicles of Fucus vesiculosus

Analysis of the gases contained in the vesicles of Fucus vesiculosusshowed that their oxygen content may be markedly enhanced byphotosynthesis. Under conditions of exposure to the air it was found that theoxygen content rarely fell much below 20 per cent. even afterprolonged periods in the dark. The carbon dioxide content of the vesicular gas varied from0 to 2·5 per cent. and appeared to bear no relationshipto metabolic activity. The results as a whole suggested that vesicle formation maypossibly be conditioned by the intensity of photosynthetic activity.     

Studies in Stomatal Behaviour: VII. EFFECTS OF ANAEROBIC CONDITIONS UPON STOMATAL MOVEMENT--A TEST OF WILLIAMS'S HYPOTHESIS OF STOMATAL MECHANISM

An experiment was carried out to investigate stomatal responsesin wheat to four ‘closing treatments’, viz. highcarbon dioxide concentration, darkness, dry air and nil, eachgiven under both aerobic and anaerobic conditions. Thus theeffect of lack of oxygen on the closing (or opening) tendencywas estimated. Changes in calculated from resistance porometer readings were used as data and reasonsare given for thinking this is the best available measure forinvestigating stomatal dynamics in wheat. Williams's hypothesisdemands that lack of oxygen should cause stomatal opening orprevent closure; the present experiment shows that anaerobicconditions significantly increase the closing tendency when‘closing treatments’ are first applied. There isalso some suggestion that oxygen-lack itself tends to causeclosure in the absence of any other ‘closing treatment’.Williams's hypothesis in its original form is thus disproved(for wheat) but the present results would be consistent withan ‘active’ uptake of water by the guard cells contributingto stomatal opening. A nearly significant interaction betweencarbon dioxide and oxygen suggests that under anaerobic conditionsa ‘closing substance’ may perhaps be formed, forexample, by the union of some intermediate in glycolysis withcarbon dioxide.     

Guard cell responses to potassium ferricyanide

Micromolar concentrations of potassium ferricyanide inhibit light-induced stomatal opening. The extent of the inhibition is dependent on the presence of carbon dioxide and the concentration of potassium ferricyanide needed to obtain 50% inhibition of stomatal opening is 40-fold higher in CO₂-free air than in normal air. The fungal toxin, fusicoccin (1 μ M ), overcame the ferricyanide inhibition of stomatal opening indicating that the electron acceptor may interact more or less directly with the activity of the plasma membrane H⁺-ATPase. Although potassium ferricyanide strongly inhibited stomatal opening, it had only minor effects on stomatal maintaining or stomatal closure due to darkness or ABA.     

Investigations of Carbon Transport in Plants: II. THE EFFECTS OF LIGHT AND DARKNESS AND SINK ACTIVITY ON TRANSLOCATION

The use of ¹¹C as a tracer has allowed repetitive measurementsof the speed of assimilate translocation to be made on singlemaize plants throughout prolonged periods of light and darkness.The speed appeared to double when the light was switched on.The time required to achieve a maximum speed, usually about3·5 cm min^–1, depended on the duration of the previousdark period. When the plant was transferred to darkness thespeed immediately decreased by about 20 per cent and continuedto decrease over the next 20 h to values of 0·5 to 0·9cm min^–1. The mean speed of translocation in tomato in the light, andother C3 plants, was usually about 1 cm min^–1. It wasreduced by 15–30 per cent when the fruit was removed orcooled from 26 to 10°C.     

Studies in the Growth of Young of Pinus sylvestris L.: I. The Annual Cycle of Assimilation and Growth

The annual cycle of growth in plants of P. sylvestris L. varyingin age from 2 to 5 years was studied mainly by following theirweight changes through periodic sampling. Immediately followingbud-break there was a period of about 1 month in which, althoughmeristematic and extension growth were rapid in both shoot androot, there was no gain in weight by the plant. There was evidencethat the needles remaining from previous years contained a considerableamount of storage material which was utilized at this time.During May and June the main increase in weight was in the newshoots, but in August and September assimilates were more evenlydivided between the new shoots on the one hand, and the oldstem and the root on the other. The average relative growth-ratesfor the months May to September were between 1·0 and0·4 per cent, per day in plants growing in favourableconditions, the lower rates being found in older plants. Duringthe months October to March slow increase in weight occurredin needles, stem, and root, and the R.G.R.s in two experimentswere 0·21 and 0·15 per cent, per day. The netassimilation rate fell with increasing age and in the summerof the fifth year was 0·06g./g./week or 0·10g./dm.²/week.This low value of N.A.R. is discussed in relation to the knownhigh rate of production of P. sylvestris stands and their evergreenhabit.     

The Kinetics of Extracellular Glycollate Production by Chlorella pyrenoidosa

Extracellular glycollate is liberated by Chlorella pyrenoidosaduring growth in medium bubbled with air or 3 per cent carbondioxide in air. With air the rate of release of glycollate percell decreases, with 3 per cent carbon dioxide it increases,with increase in cell number. Glycollate is released duringshort-term experiments when C. pyrenoidasa, grown under lowlight and high carbon dioxide, is transferred suddenly to highlight and low carbon dioxide. No other combination of thesefactors produces a comparable release of glycollate. The quantityof glycollate released in short-term experiments increases exponentiallywith the relative growth-rate of the culture from which thecells are derived. A crucial condition for maximum glycollaterelease is that growth of the culture prior to the experimentshould not be limited by carbon-dixoide concentration. The effectof pH is related to its effect on growth-rate; i.e. C. pyrenoidosahas a lower relative growth-rate at pH 8.3 and produces correspondinglyless glycollate than faster growing cultures at pH 6.4. Duringshort-term experiments under high light and low carbon dioxidethe rate of glycollate release drops after 50–100 minutessuggesting exhaustion of the glycollate precursor.     

Azetidine-2-carboxylic Acid and Lily Pollen Tube Elongation

Azetidine-2-carboxylic acid (AZC), which occurs naturally inLiliaceous plants, is reported to be a proline (pro) analoguePlant cell walls contain ‘extensin’, which is richin hydroxyproline (hyp). Peptidyl hyp arises through hydroxylationof peptidyl pro followed by glycosylation (arabinose attachment)of hyp Because AZC replaces peptidyl prolyl residues, it maybe a useful tool for evaluating the significance of hyp-o-arabinoselinkages in cell elongation. Therefore, we determined the effectof AZC on [¹⁴C]pro uptake, incorporation and conversion to wall-bound[¹⁴C]hyp in relation to elongation of lily pollen tubes whosewalls consist, in part, of hyp-containing glycopeptides TheAZC suppressed pollen germination 9–42 per cent (1–10mM) and subsequent tube elongation 40–54 per cent (0·1–1mM without affecting respiration In contrast, similar hyp concentrationswere without effect on tube elongation Whereas uptake of [¹⁴C]prowas 16·5–6·2 per cent of the control at0·1–1 mM AZC, [¹⁴C]leucine uptake was 85–25per cent of the control. Light microscope radioautography revealedfewer silver grains over tubes elongated in 0·1–1mM AZC than in its absence. Incorporation of [¹⁴C]pro into tnchloroaceticacid (TCA)-precipitable cytoplasm was reduced by only 10 percent at 0·01–1 mM but 43 per cent at 10 mM AZCGel filtration of cytoplasm from pollen germinated without AZCbut with [¹⁴C]pro resulted in labelled void volume (V) and threeretarded peaks (RI–III) Incorporation into V and RI wasinhibited at both 0·01 and 1 mM AZC These AZC concentrationsreduced conversion of [¹⁴C]pro to wall-bound hyp by 20 percent However, total incorporation of [¹⁴C]pro into salt-water-purifiedwall fractions was suppressed 47–53 per cent (0·1–1mM AZC). Lilium longiflorum, lily, hydroxyproline, proline, azetidine-2-carboxylic acid, pollen, pollen tube elongation     

In-Phase Cycling of Photosynthesis and Conductance at Saturating Carbon Dioxide Pressure Induced by Increases in Water Vapour Pressure Deficit

Bunce, J. A. 1987. In-phase cycling of photosynthesis and conductanceat saturating carbon dioxide pressure induced by increases inwater vapour pressure deficit.—J. exp. Bot. 38: 1413–1420. The leaf to air water vapour deficit was increased suddenlyfrom about 1·0 to 2·5 IcPa for single leaves ofsoybean (Glycine max L. Merr.) plants held at 30 °C, 2·0mmol m ^–2 s^–1 photosynthetic photon flux density(PPFD) and carbon dioxide pressures saturating to photosynthesis.After a lag of about 10 min, photosynthetic rate and stomatalconductance to water vapour began to decrease, and then cycledin phase with each other. The period of the cydes was about20 min. During these cycles the substomatal carbon dioxide pressurewas constant in the majority of leaves examined, and was alwaysabove saturation for photosynthesis. Epidermal impressions showedthat most stomata changed in aperture during the cycles, andthat very few were ever fully closed. Water potential measuredon excised discs changed by at most 0·1 MPa from theminima to the maxima in transpiration rate. In contrast, forleaves of sunflower (Helianthus animus L.) grown at low PPFD,the increase in VPD led to leaf wilting and decreased photosynthesis,followed by recovery of turgor and photosynthesis as stomatalconductance began to decrease. In these leaves photosynthesisand conductance then cycled approximately 180° out of phase.It is suggested that in soybeans decreased leaf conductanceinduced by high VPD provided a signal which decreased the rateof photosynthesis at carbon dioxide saturation by a mechanismthat was not related to a water deficit in the mesophyll. Key words: Photosynthesis, stomatal conductance, cycling, vapour pressure deficit     

Microbiological Synthesis of Fat: THE FORMATION OF FAT FROM SUCROSE

1. Forty-three strains of moulds, selected from ten species,were examined to ascertain their capabilities as producers offat, when grown in four different solutions of nutrient inorganicsalts together with sucrose. The three highest yields of feltwere given by Penicillium javanicum van Beyma, P. Soppi Zaleski,and Aspergillus nidulans Eidam, in that order. The fat contenton felt weight was maximal at 34·8 per cent, with P.Soppi, at 28·4 per cent. with Fusarium lini(1), and at25·8 per cent. with A. nidulans; on sugar utilized itwas maximal at 11·4 per cent. with P. Soppi, at 7·9per cent, with A. nidulans, and at 5·6 per cent. withF. lini(1). 2. Other moulds considered worthy of further trial were P. spinulosum,P. aurantiobrunneum, F. graminearum, A. flavus, F. oxysporum,A. clavatus, and P. oxalicum.     

Rates of Photosynthesis and Respiration in Relation to Stomatal Movements in Leaves Treated with {alpha}-Hydroxysulphonate and Glycollate

Leaves of tobacco and Xanthium pennsylvanicum were treated withsolutions of -hydroxysulphonate and glycollate in normal air.During the treatment with -hydroxysulphonate the intercellular-spacecarbon dioxide concentration of the leaves increased owing toa decrease in photosynthetic activity. It seems probable thatthese increases in internal carbon dioxide concentration werethe cause of stomatal closure, because when treated leaves wereflushed with carbon dioxide free air the stomata reopened. Experimentalresults also showed that the accumulation of glycollate, resultingfrom treatment with -hydroxysulphonate, could not be held responsiblefor stomatal closure because treatment with glycollate alonehad no influence on either the internal carbon dioxide concentrationor on the stomata of the leaves.     

Investigations into the Stomatal Physiology of Upland Cotton: 2. Calibration of the Infiltration Method against Leaf and Stomatal Resistances

Data are presented to show that reductions in leaf water contentof up to 12 per cent, are associated with increases of mesophyllresistance to viscous flow of air of from 50 to 130 per cent.This effect may explain the variation observed in the relationshipbetween Infiltration Score (I), and the logarithm of leaf resistance,measured porometrically, for leaves with different water contents. Highly significant linear correlations were obtained by plottingthe llogarithm of the resistance iof 10,000 stomata againstI for leaves from plants grown in the field (r = –0.92)kjor in the greenhouse (kr = –o.94). For the two se5tsof material the slopes of the loines were similar, but a differencein levels was highly significant. This is considered in relationto leaf and stomatal structure. The use of the infiltrationmethod for deriving estimates of stomatal resistance is discussed.     

THE EFFECT OF CARBON DIOXIDE TENSION ON TISSUE METABOLISM (RETINA)

The metabolism of rat retina was found to be sensitive to the concentration of the carbon dioxide-bicarbonate buffer system. Increasing the carbon dioxide from 1 per cent to 5 per cent at constant pH nearly doubled both respiration and glycolysis. Increasing the carbon dioxide at constant pH from 5 per cent to 20 per cent had no effect on glycolysis, but depressed the Q _OO2 from 31 to 19. In a medium containing glucose and the 1 per cent carbon dioxide-bicarbonate buffer, the addition of succinate increased the Q _OO2 from 12 to 26, without affecting glycolysis. In a medium containing glucose and phosphate, succinate had no significant effect.     

铁皮石斛叶片光合作用的碳代谢途径

 利用LI-6400光合测定系统测定了不同天气条件下铁皮石斛（Dendrobium officinale）叶片24h CO2吸收的动态以及CO2吸收对光强和温度的响应。晴天的白天和夜间铁皮石斛都能吸收CO2,中午CO2吸收速率为负值, CO2的交换方式具景天酸代谢途径(CAM)的特点。阴雨天,只有白天吸收CO2,夜间表现为暗呼吸,光合作用碳代谢的途径为C3途径。在多云的天气条件下,白天吸收CO2,并持续至日落后。夜间21∶00仍有CO2吸收,23∶00以后至次日凌晨处于暗呼吸状态。在500 μmol·m-2·s-1光照件下,20℃出现最大CO2吸收值。在夜间,25℃时CO2的吸收速率最高。有光和无光条件下,低温或高温引起CO2吸收速率下降均为非气孔因素所致。晴天上午,铁皮石斛叶片的表观量子产额为0.035,光合补偿点为2.9μmol·m-2·s-1,饱和光强为500μmol·m-2·s-1,强光下出现光抑制现象。叶片受到强光预先照射后,即使光照减弱光抑制效应仍保持一段时间,致使光合补偿点升高,表观量子产额下降,相同光强下的CO2吸收效率降低。结果表明：铁皮石斛为兼性CAM植物,随着环境条件的变化,其光合作用在景天酸代谢途径(CAM)与C3途径间变化。