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1.
A controlled growth chamber experiment was conducted to investigate the short-term water use and photosynthetic responses of 30-d-old carrot seedlings to the combined effects of CO2 concentration (50–1 050 μmol mol−1) and moisture deficits (−5, −30, −55, and −70 kPa). The photosynthetic response data was fitted to a non-rectangular hyperbola model. The estimated parameters were compared for effects of moisture deficit and elevated CO2 concentration (EC). The carboxylation efficiency (α) increased in response to mild moisture stress (−30 kPa) under EC when compared to the unstressed control. However, moderate (−55 kPa) and extreme (−70 kPa) moisture deficits reduced α under EC. Maximum net photosynthetic rate (P Nmax) did not differ between mild water deficit and unstressed controls under EC. Moderate and extreme moisture deficits reduced P Nmax by nearly 85 % compared to controls. Dark respiration rate (R D) showed no consistent response to moisture deficit. The CO2 compensation concentration (Γ) was 324 μmol mol−1 for −75 kPa and ranged 63–93 μmol mol−1 for other moisture regimes. Interaction between moisture deficit and EC was noticed for P N, ratio of intercellular and ambient CO2 concentration (C i/C a), stomatal conductance (g s ), and transpiration rate (E). P N was maximum and C i/C a was minimum at −30 kPa moisture deficit and at C a of 350 μmol mol−1. The g s and E showed an inverse relationship at all moisture deficit regimes and EC. Water use efficiency (WUE) increased with moisture deficit up to −55 kPa and declined thereafter. EC showed a positive influence towards sustaining P N and increasing WUE only under mild moisture stress, and no beneficial effects of EC were noticed at moderate or extreme moisture deficits.  相似文献   

2.
Net photosynthetic rate (P N) of leaves grown under free-air CO2 enriched condition (FACE, about 200 μmol mol−1 above ambient air) was significantly lower than P N of leaves grown at ambient CO2 concentration (AC) when measured at CO2 concentration of 580 μmol mol−1. This difference was found in rice plants grown at normal nitrogen supply (25 g m−2; NN-plants) but not in plants grown at low nitrogen supply (15 g m−2; LN-plants). Namely, photosynthetic acclimation to FACE was observed in NN-plants but not in LN-plants. Different from the above results measured in a period of continuous sunny days, such photosynthetic acclimation occurred in NN-plants, however, it was also observed in LN-plants when P N was measured before noon of the first sunny day after rain. Hence strong competition for the assimilatory power between nitrogen (N) and carbon (C) assimilations induced by an excessive N supply may lead to the photosynthetic acclimation to FACE in NN-plants. The hypothesis is supported by the following facts: FACE induced significant decrease in both apparent photosynthetic quantum yield (Φc) and ribulose-1,5-bisphosphate (RuBP) content in NN-plants but not in LN-plants.  相似文献   

3.
We determined the interactive effects of irradiance, elevated CO2 concentration (EC), and temperature in carrot (Daucus carota var. sativus). Plants of the cv. Red Core Chantenay (RCC) were grown in a controlled environmental plant growth room and exposed to 3 levels of photosynthetically active radiation (PAR) (400, 800, 1 200 μmol m−2 s−1), 3 leaf chamber temperatures (15, 20, 30 °C), and 2 external CO2 concentrations (C a), AC and EC (350 and 750 μmol mol−1, respectively). Rates of net photosynthesis (P N) and transpiration (E) and stomatal conductance (g s ) were measured, along with water use efficiency (WUE) and ratio of internal and external CO2 concentrations (C i/C a). P N revealed an interactive effect between PAR and C a. As PAR increased so did P N under both C a regimes. The g s showed no interactive effects between the three parameters but had singular effects of temperature and PAR. E was strongly influenced by the combination of PAR and temperature. WUE was interactively affected by all three parameters. Maximum WUE occurred at 15 °C and 1 200 μmol m−2 s− 1 PAR under EC. The C i /C a was influenced independently by temperature and C a. Hence photosynthetic responses are interactively affected by changes in irradiance, external CO2 concentration, and temperature. EC significantly compensates the inhibitory effects of high temperature and irradiance on P N and WUE.  相似文献   

4.
The effect of elevated carbon dioxide concentration on the changes in the biomass, photosynthesis and nutrient composition was investigated in two leafy vegetables. Spinach (Spinacia oleracea L.) and fenugreek (Trigonella foenum-graecum L.) plants were grown in open top chambers under either ambient (ACO2, 350 ± 50 μmol mol−1) or elevated (ECO2, 600 ± 50 μmol mol−1) CO2 concentration and analyzed 40, 60 and 80 days after exposure. The plants grown in ECO2 had higher net photosynthetic rate and lower stomatal conductance when compared with the plants grown in ACO2. ECO2 also changed the nutrient composition: a lower N, Mg and Fe contents and higher C and Ca contents were observed in the leaves of plants exposed to ECO2 than in those grown at ACO2.  相似文献   

5.
Chaetoceros convolutus and C. concavicornis have been implicated in the death of salmon in netpens in the Pacific Northwest by damaging the salmon's gills. To better understand how environmental factors affect the distribution of these two species, the interacting effects of light, temperature and salinity on growth rate were examined by growing these species under a range of temperatures (4–18 °C), light (10–175 μmol photon m−2 s−1) and salinities (10–30‰). For C. convolutus, the growth rate showed a hyperbolic relationship with irradiance at 8, 14 and 18 °C and light saturation occurred at 9, 14 and 20 μmol photon mt s−1 respectively. At 4 °C for C. convolutus and 8 °C for C. concavicornis, cells grew at μmax, even at the lowest irradiances tested (10 μmol photon m−2 s−1). For C. convolutus, the amount of light required to saturate growth rate increased with temperature in an approximately linear fashion. The Q10 was 1.88, calculated by averaging over both species. C. concavicornis was the more euryhaline species growing at salinities as low as 17.5‰, while C. convolutus grew only at 25‰ and above.  相似文献   

6.
The effects of salinity (sea water at 0 ‰ versus 30 ‰) on gross rates of O2 evolution (J O2) and net rates of CO2 uptake (P N) were measured in the halotolerant estuarine C4 grasses Spartina patens, S. alterniflora, S. densiflora, and Distichlis spicata in controlled growth environments. Under high irradiance, salinity had no significant effect on the intercellular to ambient CO2 concentration ratio (C i/C a). However, during photosynthesis under limiting irradiance, the maximum quantum efficiency of CO2 fixation decreased under salinity across species, suggesting there is increased leakage of the CO2 delivered to the bundle sheath cells by the C4 pump. Growth under salinity did not affect the maximum intrinsic efficiency of photosystem 2, PS2 (FV/FM) in these species, suggesting salinity had no effect on photosynthesis by inactivation of PS2 reaction centers. Under saline conditions and high irradiance, P N was reduced by 75 % in Spartina patens and S. alterniflora, whereas salinity had no effect on P N in S. densiflora or D. spicata. This inhibition of P N in S. patens and S. alterniflora was not due to an effect on stomatal conductance since the ratio of C i/C a did not decrease under saline conditions. In growth with and without salt, P N was saturated at ∼500 μmol(quantum) m−2 s−1 while J O2 continued to increase up to full sunlight, indicating that carbon assimilation was not tightly coupled to photochemistry in these halophytic species. This increase in alternative electron flow under high irradiance might be an inherent function in these halophytes for dissipating excess energy.  相似文献   

7.
The effect of elevated CO2 concentration (CE) on leaf chlorophyll (Chl) and nitrogen (N) contents and photosynthetic rate (PN) was evaluated during the post-flowering stages of rice grown at CE (570 ± 50 μmol mol−1) in open top chamber (OTC), at ambient CO2 concentration (∼ 365 μmol mol−1) in OTC and at open field. Thirty-five day old seedlings were transplanted in OTCs or in field and allowed to grow till maturity. Chl and N contents were highest at the time of flowering and thereafter it started to decline. The rate of decline in Chl and N contents was faster in plants grown under CE mostly in later part of growth. Irrespective of treatment difference, flag leaf contained the highest amount of Chl and N than penultimate and third leaf. The higher PN was observed in leaves under CE than in the leaves in other two growing conditions. Considering growth stage, PN was the highest at flowering which reduced at the later part of growth due to degradation of Chl and N content of the leaf. Under CE it was 40.02 μmol m−2 s−1 at flowering and it reduced to only 14.77 μmol m−2 s−1 at maturity stage. The beneficial effect of CE in increasing leaf PN may be maintained by applying extra dose of nitrogen at the later stages of plant growth.  相似文献   

8.
The effect of drought stress (DS) on photosynthesis and photosynthesis-related enzyme activities was investigated in F. pringlei (C3), F. floridana (C3–C4), F. brownii (C4-like), and F. trinervia (C4) species. Stomatal closure was observed in all species, probably being the main cause for the decline in photosynthesis in the C3 species under ambient conditions. In vitro ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and stromal fructose 1,6-bisphosphatase (sFBP) activities were sufficient to interpret the net photosynthetic rates (P N), but, from the decreases in P N values under high CO2 (C a = 700 μmol mol− 1) it is concluded that a decrease in the in vivo rate of the RuBPCO reaction may be an additional limiting factor under DS in the C3 species. The observed decline in the photosynthesis capacity of the C3–C4 species is suggested to be associated both to in vivo decreases of RuBPCO activity and of the RuBP regeneration rate. The decline of the maximum P N observed in the C4-like species under DS was probably attributed to a decrease in maximum RuBPCO activity and/or to decrease of enzyme substrate (RuBP or PEP) regeneration rates. In the C4 species, the decline of both in vivo photosynthesis and photosynthetic capacity could be due to in vivo inhibition of the phosphoenolpyruvate carboxylase (PEPC) by a twofold increase of the malate concentration observed in mesophyll cell extracts from DS plants.  相似文献   

9.
Effects of three levels of photosynthetic photon flux (PPF: 60, 160 and 300 μmol m−2s−1) were investigated in one-month-old Phalaenopsis plantlets acclimatised ex vitro. Optimal growth, chlorophyll and carotenoid concentations, and a high carotenoid:chlorophyll a ratio were obtained at 160 μmol m−2s−1, while net CO2 assimilation (A), stomatal conductance (g), transpiration rate (E) and leaf temperature peaked at 300 μmol m−2s−1, indicating the ability of the plants to grow ex vitro. Adverse effects of the highest PPF were reflected in loss of chlorophyll, biomass, non-protein thiol and cysteine, but increased proline. After acclimatisation, glucose-6-phosphate dehydrogenase, shikimate dehydrogenase, phenylalanine ammonia-lyase (PAL) and cinnamyl alcohol dehydrogenase (CAD) increased, as did lignin. Peroxidases (POD), which play an important role in lignin synthesis, were induced in acclimatised plants. Polyphenol oxidase (PPO) and β-glucosidase (β-GS) activities increased to a maximum in acclimatised plants at 300 μmol m−2s−1. A positive correlation between PAL, CAD activity and lignin concentration was observed, especially at 160 and 300 μmol m−2s−1. The study concludes that enhancement of lignin biosynthesis probably not only adds rigidity to plant cell walls but also induces defence against radiation stress. A PPF of 160 μmol m−2s−1was suitable for acclimatisation when plants were transferred from in vitro conditions.  相似文献   

10.
Quercus ilex plants grown on two different substrates, sand soil (C) and compost (CG), were exposed to photosynthetic photon flux densities (PPFD) at 390 and 800 μmol(CO2) mol−1 (C390 and C800). At C800 both C and CG plants showed a significant increase of net photosynthetic rate (P N) and electron transport rate (ETR) in response to PPFD increase as compared to C390. In addition, at C800 lower non-photochemical quenching (NPQ) values were observed. The differences between C390 and C800 were related to PPFD. The higher P N and ETR and the lower dissipative processes found in CG plants at both CO2 concentrations as compared to C plants suggest that substrate influences significantly photosynthetic response of Q. ilex plants. Moreover, short-term exposures at elevated CO2 decreased nitrate photo-assimilation in leaves independently from substrate of growth.  相似文献   

11.
Citrus volkameriana (L.) plants were grown for 43 d in nutrient solutions containing 0, 2, 14, 98, or 686 μM Mn (Mn0, Mn2, Mn14, Mn98, and Mn686, respectively). To adequately investigate the combined effects of Mn nutrition and irradiance on photosystem 2 (PS2) activity, irradiance response curves for electron transport rate (ETR), nonphotochemical quenching (qN), photochemical quenching (qP), and real photochemical efficiency of PS2 (ΦPS2) were recorded under 10 different irradiances (66, 96, 136, 226, 336, 536, 811, 1 211, 1 911, and 3 111 μmol m−2 s−1, I66 to I3111, respectively) generated with the PAM-2000 fluorometer. Leaf chlorophyll content was significantly lower under Mn excess (Mn686) compared to Mn0; its highest values were recorded in the treatments Mn2-Mn98. However, ETR and ΦPS2 values were significantly lower under Mn0 compared to the other Mn treatments, when plants were exposed to irradiances ≥96 μmol m−2 s−1. Furthermore, Mn0 plants had significantly higher values of qN and lower values of qP at irradiances ≤226 and ≥336 μmol m−2 s−1, respectively, than those grown under Mn2-Mn686. Irrespective of Mn treatment, the values of ΦPS2 and qN decreased, while those of qP increased progressively by increasing irradiance from I136 to I3111. Finally, Mn2-Mn98 plants were less sensitive to photoinhibition of photosynthesis (≥811 μmol m−2 s−1) than the Mn686 (≥536 μmol m−2 s−1) and Mn0 (≥336 μmol m−2 s−1) ones.  相似文献   

12.
Photoautotrophic micropropagation of Spathiphyllum   总被引:2,自引:0,他引:2  
In order to maximize yield, Spathiphyllum, an ornamental plant, was cultured in vitro in novel culture vessels termed Vitron. The best growth was obtained by culturing plantlets on sugar-free liquid medium under CO2 enrichment (3 000 μmol mol−1 24 h−1 d−1) at a low photon flux density (PPFD of 45 μmol m−2 s−1), suggesting that the novel Vitron culture system is suitable for the photoautotrophic micropropagation of Spathiphyllum.  相似文献   

13.
To elucidate whether dipterocarp species, dominant late-successional species of tropical forests in Southeast Asia, actually have a disadvantage when planted on open site in terms of their photosynthetic characteristics, we investigated photosynthesis in dipterocarp seedlings planted in the open on degraded sandy soils in southern Thailand. These species were compared with seedlings of Acacia mangium Willd., a fast-growing tropical leguminous tree, which is often planted on degraded open site in Southeast Asia. The dipterocarp seedlings had an irradiance-saturated net photosynthetic rate (P N), stomatal conductance (g s), carboxylation efficiency, and photosynthetic capacity comparable to or superior to those of A. mangium. In particular, seedlings of Dipterocarpus obtusifolius Teijsm. ex Miq. showed an irradian-ce-saturated P N of 21 μmol m−2 s−1, a value higher than any previously reported for a dipterocarp species, accompanied by high g s (0.7 mol m−2 s−1) and high photosynthetic capacity. Thus dipterocarp species do not necessarily have a disadvantage in terms of their photosynthetic characteristics on open sites with degraded sandy soils.  相似文献   

14.
Two cultivars (Katy and Erhuacao) of apricot (Prunus armeniaca L.) were evaluated under open-field and solar-heated greenhouse conditions in northwest China, to determine the effect of photosynthetic photon flux density (PPFD), leaf temperature, and CO2 concentration on the net photosynthetic rate (P N). In greenhouse, Katy registered 28.3 μmol m−2 s−1 for compensation irradiance and 823 μmol m−2 s−1 for saturation irradiance, which were 73 and 117 % of those required by Erhuacao, respectively. The optimum temperatures for cvs. Katy and Erhuacao were 25 and 35 °C in open-field and 22 and 30 °C in greenhouse, respectively. At optimal temperatures, P N of the field-grown Katy was 16.5 μmol m−2 s−1, 21 % less than for a greenhouse-grown apricot. Both cultivars responded positively to CO2 concentrations below the CO2 saturation concentration, whereas Katy exhibited greater P N (18 %) and higher carboxylation efficiency (91 %) than Erhuacao at optimal CO2 concentration. Both cultivars exhibited greater photosynthesis in solar-heated greenhouses than in open-field, but Katy performed better than Erhuacao under greenhouse conditions.  相似文献   

15.
Čajánek  M.  Hudcová  M.  Kalina  J.  Lachetová  I.  Špunda  V. 《Photosynthetica》1999,37(2):295-306
Effects of short-term exposure to different irradiances on the function of photosystem 2 (PS2) were studied for barley grown at low (LI; 50 μmol m−2 s−1) and high (HI; 1 100 μmol m−2 s−1) irradiances. HI barley revealed higher ability to down-regulate the light-harvesting within PS2 after exposure to high irradiance as compared to LI plants. This ability was estimated from the light-induced decreases of F685/F742 and E476/E436 in emission and excitation spectra of 77 K chlorophyll (Chl) a fluorescence in vivo which was 65 and 10 % for HI plants as compared to 30 and 2 % for LI plants, respectively. For LI plants this protective down-regulation of the light-harvesting of PS2 was saturated at 430 μmol m−2 s−1, and progressive PS2 photodamage was induced at higher irradiances. After exposure of LI segments to 2 200 μmol m−2 s−1 a pronounced maximum at 700 nm appeared in emission spectrum of 77 K Chl a fluorescence. Based on complementary analysis of 77 K excitation spectra measured at the emission wavelength 685 nm we suggest that this emission maximum may be attributed to the formation of aggregates of light-harvesting complexes of PS2 (LHC2) with part of PS2 core during progressive PS2 photodamage. Our results can be explained assuming different contributions of LHC2 and PS2 core to the total nonradiative dissipation of absorbed excitation energy for the LI and HI barley. This revised version was published online in August 2006 with corrections to the Cover Date.  相似文献   

16.
In sunflower (Helianthus annuus L.) grown under controlled conditions and subjected to drought by withholding watering, net photosynthetic rate (P N) and stomatal conductance (g s) of attached leaves decreased as leaf water potential (Ψw) declined from −0.3 to −2.9 MPa. Although g s decreased over the whole range of Ψw, nearly constant values in the intercellular CO2 concentrations (C i) were observed as Ψw decreased to −1.8 MPa, but C i increased as Ψw decreased further. Relative quantum yield, photochemical quenching, and the apparent quantum yield of photosynthesis decreased with water deficit, whereas non-photochemical quenching (qNP) increased progressively. A highly significant negative relationship between qNP and ATP content was observed. Water deficit did not alter the pyridine nucleotide concentration but decreased ATP content suggesting metabolic impairment. At a photon flux density of 550 μmol m−2 s−1, the allocation of electrons from photosystem (PS) 2 to O2 reduction was increased by 51 %, while the allocation to CO2 assimilation was diminished by 32 %, as Ψw declined from −0.3 to −2.9 MPa. A significant linear relationship between mean P N and the rate of total linear electron transport was observed in well watered plants, the correlation becoming curvilinear when water deficit increased. The maximum quantum yield of PS2 was not affected by water deficit, whereas qP declined only at very severe stress and the excess photon energy was dissipated by increasing qNP indicating that a greater proportion of the energy was thermally dissipated. This accounted for the apparent down-regulation of PS2 and supported the protective role of qNP against photoinhibition in sunflower.  相似文献   

17.
Summary An efficient protocol for in vitro propagation of an aromatic and medicinal herb Ocimum basilicum L. (sweet basil) through axillary shoot proliferation from nodal explants, collected from field-grown plants, is described. High frequency bud break and maximum number of axillary shoot formation was induced in the nodal explants on Murashige and Skoog (1962) medium (MS) containing N6-benzyladenine (BA). The nodal explants required the presence of BA at a higher concentration (1.0 mg·l−1, 4.4 μM) at the initial stage of bud break; however, further growth and proliferation required transfer to a medium containing BA at a relatively low concentration (0.25 mg·gl−1, 1.1 μM). Gibberellic (GA3) at 0.4 mg·l−1 (1.2 μM) added to the medium along with BA (1.0 mg·l−1, 4.4 μM) markedly enhanced the frequency of bud break. The shoot clumps that were maintained on the proliferating medium for longer durations, developed inflorescences and flowered in vitro. The shoots formed in vitro were rooted on half-strength MS supplemented with 1.0 mg·l−1 (5.0 μM) indole-3-butyric acid (IBA). Rooted plantlets were successfully acclimated in vermi-compost inside a growth chamber and eventually established in soil. All regenerated plants were identical to the donor plants with respect to vegetative and floral morphology.  相似文献   

18.
Six months old in vitro-grown Anoectochilus formosanus plantlets were transferred to ex-vitro acclimation under low irradiance, LI [60 μmol(photon) m−2 s−1], intermediate irradiance, II [180 μmol(photon) m−2 s−1], and high irradiance, HI [300 μmol(photon) m−2 s−1] for 30 d. Imposition of II led to a significant increase of chlorophyll (Chl) b content, rates of net photosynthesis (P N) and transpiration (E), stomatal conductance (g s), electron transfer rate (ETR), quantum yield of electron transport from water through photosystem 2 (ΦPS2), and activity of ribulose-1,5-bisphosphate carboxylase/ oxygenase (RuBPCO, EC 4.1.1.39). This indicates that Anoectochilus was better acclimated at II compared to LI treatment. On the other hand, HI acclimation led to a significant reduction of Chl a and b, P N, E, g s, photochemical quenching, dark-adapted quantum efficiency of open PS2 centres (Fv/Fm), probability of an absorbed photon reaching an open PS2 reaction centre (Fv′/Fm′), ETR, ΦPS2, and energy efficiency of CO2 fixation (ΦCO2PS2). This indicates that HI treatment considerably exceeded the photo-protective capacity and Anoectochilus suffered HI induced damage to the photosynthetic apparatus. Imposition of HI significantly increased the contents of antheraxanthin and zeaxanthin (ZEA), non-photochemical quenching, and conversion of violaxanthin to ZEA. Thus Anoectochilus modifies its system to dissipate excess excitation energy and to protect the photosynthetic machinery.  相似文献   

19.
Z.-P. Ye 《Photosynthetica》2007,45(4):637-640
The calculated maximum net photosynthetic rate (P N) at saturation irradiance (I m) of 1 314.13 μmol m−2 s−1 was 25.49 μmol(CO2) m−2 s−1, and intrinsic quantum yield at zero irradiance was 0.103. The results fitted by nonrectangular hyperbolic model, rectangular hyperbolic method, binomial regression method, and the new model were compared. The maximum P N values calculated by nonrectangular hyperbolic model and rectangular hyperbolic model were higher than the measured values, and the I m calculated by nonrectangular hyperbolic model and rectangular hyperbolic model were less than measured values. Results fitted by new model showed that the response curve of P N to I was nonlinear at low I for Oryza sativa, P N increased nonlinearly with I below saturation value. Above this value, P N decreased nonlinearly with I.  相似文献   

20.
Photosynthetic rates of green leaves (GL) and green flower petals (GFP) of the CAM plant Dendrobium cv. Burana Jade and their sensitivities to different growth irradiances were studied in shade-grown plants over a period of 4 weeks. Maximal photosynthetic O2 evolution rates and CAM acidities [dawn/dusk fluctuations in titratable acidity] were higher in leaves exposed to intermediate sunlight [a maximal photosynthetic photon flux density (PPFD) of 500–600 μmol m−2 s−1] than in leaves grown under full sunlight (a maximal PPFD of 1 000–1 200 μmol m−2 s−1) and shade (a maximal PPFD of 200–250 μmol m−2 s−1). However, these two parameters of GFP were highest in plants grown under the shade and lowest in full sun-grown plants. Both GL and GFP of plants exposed to full sunlight had lower predawn Fv/Fm [dark adapted ratio of variable to maximal fluorescence (the maximal photosystem 2 yield without actinic irradiation)] than those of shade-grown plants. When exposed to intermediate sunlight, however, there were no significant changes in predawn Fv/Fm in GL whereas a significant decrease in predawn Fv/Fm was found in GFP of the same plant. GFP exposed to full sunlight exhibited a greater decrease in predawn Fv/Fm compared to those exposed to intermediate sunlight. The patterns of changes in total chlorophyll (Chl) content of GL and GFP were similar to those of Fv/Fm. Although midday Fv/Fm fluctuated with prevailing irradiance, changes of midday Fv/Fm after exposure to different growth irradiances were similar to those of predawn Fv/Fm in both GL and GFP. The decreases in predawn and midday Fv/Fm were much more pronounced in GFP than in GL under full sunlight, indicating greater sensitivity in GFP to high irradiance (HI). In the laboratory, electron transport rate and photochemical and non-photochemical quenching of Chl fluorescence were also determined under different irradiances. All results indicated that GFP are more susceptible to HI than GL. Although the GFP of Dendrobium cv. Burana Jade require a lower amount of radiant energy for photosynthesis and this plant is usually grown in the shade, is not necessarily a shade plant.  相似文献   

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