Light and temperature effects on the growth rate of three freshwater [2pt] algae isolated from a eutrophic lake

Effects of light and temperature, on the growth of three freshwater green algae isolated from an eutrophic lake and identified as Selenastrum minutum, Coelastrum microporum f. astroidea and Cosmarium subprotumidumwere studied in batch cultures under non-nutrient limited conditions. Experiments were performed to determine the growth rate over a wide range of light intensities (30–456 mol m^–2 s^–1) and temperature (15–35°C), using a 15/9 (light/dark) photoperiod cycle. The maximum growth rates and the optimum light intensities at a temperature of 35°C were 1.73 d^–1 and 420 mol m^–2 s^–1for Selenastrum minutum, 1.64 d^–1 and 400 mol m^–2 s^–1 for Coelastrum microporum and 1.00 d^–1 and 400 mol m^–2 s¹ for Cosmarium subprotumidum. The results were fitted with the mathematical models of Steele (1965), Platt & Jassby (1976) and Peeters & Eilers (1978). Steele's function and equation of Platt & Jassby don't describe correctly the relationship between the growth and light intensity. In the opposite, the equation of Peeters & Eilers provides the best fit for the three species.     

Growth and phosphate uptake kinetics of the cyanobacterium,Cylindrospermopsis raciborskii (Cyanophyceae) in throughflow cultures

• 1 Cylindrospermopsis raciborskii occupies a rapidly expanding geographical area. Its invasive success challenges eutrophication control in many lakes. To understand better the load‐dependent behaviour of this nitrogen fixing cyanobacterium under in situ conditions, we studied P‐dependent growth of a C. raciborskii strain under continuous and pulsed P supply.
• 2 The Droop model reasonably described P‐dependent growth in the continuously supplied chemostats. Large P pulses, however, caused a delay in growth and cells subject to P pulses grew slower than their counterparts with the same P quota supplied continuously.
• 3 The kinetics of P uptake indicated that C. raciborskii is opportunistic with respect to P. Its high excess P storage capacity after a saturating P pulse (Q_ex=95 µg P [mg C]^‐1) and P‐specific uptake capacity (U_max = V_max/Q_P=150–1200) are indicative of storage adaptation. At the same time, the affinity of the P uptake system (U_max/K = 800–4000) is also high.
• 4 Rate of leakage exceeded that of the steady state net P uptake by one to two orders of magnitude. Growth affinity of C. raciborskii (µ_max/K_µ≈ 20) was relatively low, presumably due to the substantial leakage.
• 5 The dynamics of the particular water body determine which trait contributes most to competitive success of C. raciborskii. In deep lakes with vertical nutrient gradients, the cyanobacterium may rely primarily on its high P storage capacity, which is coupled to a lack of short‐term feedback inhibition and efficient buoyancy regulation. In lakes without such gradients, high P uptake affinity may be vitally important.

     

Effect of irradiance and temperature on the photosynthesis of a cultivated red alga,Pyropia tenera (= Porphyra tenera), at the southern limit of distribution in Japan

The effect of irradiance and temperature on the photosynthesis of the red alga, Pyropia tenera, was determined for maricultured gametophytes and sporophytes collected from a region that is known as one of the southern limits of its distribution in Japan. Macroscopic gametophytes were examined using both pulse‐amplitude modulated fluorometry and/or dissolved oxygen sensors. A model of the net photosynthesis–irradiance (P‐E) relationship of the gametophytes at 12°C revealed that the net photosynthetic rate quickly increased at irradiances below the estimated saturation irradiance of 46 μmol photons m^?2 s^?1, and the compensation irradiance was 9 μmol photons m^?2 s^?1. Gross photosynthesis and dark respiration for the gametophytes were also determined over a range of temperatures (8–34°C), revealing that the gross photosynthetic rates of 46.3 μmol O₂ mg_chl‐a^?1 min^?1 was highest at 9.3 (95% Bayesian credible interval (BCI): 2.3–14.5)°C, and the dark respiration rate increased at a rate of 0.93 μmol O₂ mg_chl‐a^?1 min^?1°C^?1. The measured dark respiration rates ranged from ?0.06 μmol O₂ mg_chl‐a^?1 min^?1 at 6°C to ?25.2 μmol O₂ mg_chl‐a^?1 min^?1 at 34°C. The highest value of the maximum quantum yield (Fv/Fm) for the gametophytes occurred at 22.4 (BCI: 21.5–23.3) °C and was 0.48 (BCI: 0.475–0.486), although those of the sporophyte occurred at 12.9 (BCI: 7.4–15.1) °C and was 0.52 (BCI: 0.506–0.544). This species may be considered well‐adapted to the current range of seawater temperatures in this region. However, since the gametophytes have such a low temperature requirement, they are most likely close to their tolerable temperatures in the natural environment.     

Snow algae of the Windmill Islands,continental Antarctica. 2. Chloromonas rubroleosa sp. nov. (Volvocales,Chlorophyta), an alga of red snow

A new, cryophilic species of the Chlamydomonadaceae, Chloromonas rubroleosa, is described from continental Antarctica. The species is characterised by a bimamillate papilla, four contractile vacuoles, numerous small discoid chloroplasts and an abundance of red-pigmented oil droplets. The local distribution of the species and its habitat conditions are described. The cells are susceptible to temperatures above 10°C and, unexpectedly, to subfreezing temperatures. The species has been isolated into pure culture and is currently maintained at the Australian Antarctic Division.     

The influence of irradiance,photoperiod and temperature on the growth kinetics of three planktonic diatoms

The influence of irradiance, photoperiod and temperature was determined for the growth kinetics of the diatoms Aulacoseira subarctica, Stephanodiscus astraea and Stephanodiscus hantzschii and the results compared with those of cyanobacteria. Irradiance and photoperiod relationships were qualitatively similar to those for cyanobacteria in that: (1) growth rate (K) was proportionally greater under short photoperiods, with ratios of K under continuous light to K under 3:21 light:dark (LD) cycles of 1·50, 1·80 and 2·96 for A. subarctica, S. astraea and S. hantzschii respectively; (2) at subsaturating irradiances, K was proportional to irradiance and independent of temperature with a negligible predicted maintenance growth rate requirement. Apparent growth efficiencies (GE) at subsaturating irradiances were 0·26±0·03, 0·42±0·03 and 0·50±0·03 divisions mol^-1m² for A. subarctica, S. astraea and S. hantzschii with the values for Stephanodiscus species comparable to values for Oscillatoria species. Under a 3:21 LD cycle at 4 °C, light-saturated growth rates were 0·066±0·004, 0·197±0·033 and 0·285±0·018 divisions day^-1 for A. subarctica, S. astraea and S. hantzschii. S. hantzschii growth rate at 4 °C exceeded maximum Oscillatoria growth rates at 23 °C and the S. astraea growth rate at 4 °C was equivalent to O. agardhii growth rate at 20 °C. Temperature increases above 4 °C gave Q₁₀ values between 4 °C and 12 °C of 3·68, 2·39 and 1·92 for A. subarctica, S. astraea and S. hantzschii, but higher temperatures resulted in minor increases in K. S. astraea growth rate peaked at 16 °C, declining sharply at higher temperatures. February to March in situ growth rates in Lough Neagh, mean temperature 4·3 °C, showed that the A. subarctica in situ K of 0·058 divisions day^-1 was close to the laboratory K at 4 °C, but that S. astraea in situ K of 0·101 divisions day^-1 was lower than the laboratory K at 4 °C.     

Isolation,Growth, Ultrastructure,and Metal Tolerance of the Green Alga,Chlamydomonas acidophila (Chlorophyta)

An acidophilic volvocine flagellate, Chlamydomonas acidophila (Volvocales) that was isolated from an acid lake, Katanuma, in Miyagi prefecture, Japan was studied for growth, ultrastructural characterization, and metal tolerance.

Chlamydomonas acidophila is obligately photoautotrophic, and did not grow in the cultures containing acetate or citrate even in the light. The optimum pH for growth was 3.5-4.5. To characterize metal tolerance, the toxic effects of Cd, Co, Cu, and Zn on this alga were also studied. Effective metal concentrations, which limited the growth by 50%, EC₅₀ were measured, after 72h of static exposure. EC₅₀s were 14.4 μM Cd²⁺, 81.3 μM Co²⁺, 141μM Cu²⁺, and 1.16 mM Zn²⁺ for 72 h of exposure. Thus, this alga had stronger tolerance to these metals than other species in the genus Chlamydomonas.     

蓝光对水稻幼苗生长及内源激素水平的影响

研究表明：与白光对照相比,蓝光明显抑制水稻幼苗的生长,并使幼苗体内的自由态IAA、GA1、玉米素和二氢玉米素含量下降,ABA含量和乙烯释放量则明显增加。说明蓝光对水稻幼苗生长的影响与其体内激素状况有关。     

不同遮阴环境下木奶果幼苗生长与生理生化的响应

木奶果是一种在幼苗期和成熟期对光适应性不同的热带乔木树种。研究不同遮阴环境下木奶果幼苗的生长及生理响应机制,以明确木奶果幼苗生长的适宜光照环境范围,对木奶果的苗木栽培及生态应用意义重大。本试验设置了5个处理,即CK(无遮阴),S3(遮光率45.3%)、S4(遮光率69.2%)、S6(遮光率80.0%)和S8(遮光率90.2%),观测停扣山和谢鞋山种源的木奶果幼苗4个月后的生长生理变化情况。结果显示:随着遮阴强度的加大,木奶果幼苗的叶面积与光合色素含量一直上升;叶绿素a/b值和过氧化氢酶(CAT)活性一直下降;丙二醛含量在S6前先呈下降趋势,而后稍有上升;株高和地径增长量先升后降,相对电导率、脯氨酸(Pro)与可溶性糖(Ss)含量则表现为先降后升。强光环境下,木奶果幼苗通过增加类胡萝卜素相对含量与CAT活性,以减缓膜脂过氧化损害;弱光环境下,木奶果幼苗通过增加叶面积与光合色素含量提高光能利用率。Pro与Ss则作为综合调节物,调节渗透压,以减轻强光与弱光带来的细胞膜胁迫压力。其中,停扣山与谢鞋山种源分别在S6与S4处理下Pro与Ss含量达最低值且株高与地径增长量达最高值。表明幼苗期的木奶果...     

Evaluation of physiological responses and tolerance to low-temperature stress of four Iceland poppy (Papaver nudicaule) varieties

Iceland poppy (Papaver nudicaule) is a boreal flowering plant and native to subpolar regions of Europe, Asia and North America, and the mountains of Central Asia. To investigate response and tolerance to low-temperature stress, the four varieties of Iceland poppy, Champagne Bubbles (CB), Domestic Variety (DV), Wonderland (WL), and Garden Gnome (GG), were subjected to low-temperature treatment from 0°C to ?9°C. The relative electrical conductivities in four varieties increased along with the decrease in temperature. Treated by ?3°C, the morphological changes of seedlings at different time points were investigated. Four varieties were differentially injured by freezing temperature. The dynamic accumulations of malondialdehyde (MDA), free proline (FP), soluble proteins (SP), and activities of superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD) during low-temperature treatment were measured and compared. A minor change in SP content during treatment was found, whereas MDA and FP were significantly induced. Complex patterns of activity alternations of SOD, POD, and CAT were found. Based on the calculation of semi-lethal temperature and comprehensive physiological parameters investigation, the tolerance of four varieties was evaluated as CB?>?GG and DV?>?WL.     

Photosynthetic responses to photosynthetically active radiation and temperature including chilling‐light stress on the heteromorphic life history stages of a brown alga,Cladosiphon okamuranus (Chordariaceae) from Ryukyu Islands,Japan

Photosynthetic responses to temperature and photosynthetically active radiation (PAR) were investigated on the heteromorphic life history stages (macroscopic and microscopic stages) of an edible Japanese brown alga, Cladosiphon okamuranus from the Ryukyu Islands. Measurements were carried out by using optical dissolved oxygen sensors and a pulse‐amplitude modulated fluorometer. Maximum net photosynthetic rates and other parameters of the Photosynthesis – PAR curves at 28°C were somewhat similar in both life history stages, without characteristic photoinhibition at 1000 μmol photons m^?2 s^?1. Results of oxygenic gross photosynthesis and dark respiration experiments over a temperature range of 8–40°C revealed similar temperature optima for both stages (29.7°C, macroscopic stage; 30.3°C, microscopic stage), which support their observed occurrences in the habitat during summer. Maximum quantum yields of photosystem II (PSII ) (F _v /F _m ) were relatively stable at low temperatures with the highest at 15.1°C for the macroscopic stage and at 16.5°C for the microscopic stage; but dropped at higher temperatures especially above 28°C. Continuous exposures (6 h) to 200 and 1000 μmol photons m^?2 s^?1 at 8, 16, and 28°C revealed greater depressions in effective quantum yields of PSII (Φ _PSII ) of the microscopic stage at 8°C, as well as its F _v /F _m that barely increased after 6 h of dark acclimation. Whereas post‐dark acclimation F _v /F _m of both stages exposed to low PAR fairly recovered at 28°C, suggesting their photosynthetic tolerance to such high temperature. Under natural conditions, both heteromorphic stages of C. okamuranus may persist throughout the year in this region. Beyond its northern limit of distribution, the microscopic stage of this species may suffer from photodamage, as enhanced by low winter temperatures; hence, its restricted occurrence.     

Effects of temperature, irradiance and photoperiod on growth and pigment content in some freshwater red algae in culture

The responses of relative growth rate (% day‐1) and pigment content (chlorophyll a, phycocyanin and phycoerythrin) to temperature, irradiance and photoperiod were analyzed in culture in seven freshwater red algae: Audouinella hermannii (Roth) Duby, Audouinella pygmaea (Kützing) Weber‐van Bosse, Batrachospermum ambiguum Montagne, Batrachospermum delicatulum (Skuja) Necchi et Entwisle,‘Chantransia’ stages of B. delicatulum and Batrachospermum macrosporum Montagne and Compsopogon coeruleus (C. Agardh) Montagne. Experimental conditions included temperatures of 10, 15, 20 and 25°C and low and high irradiances (65 and 300 μmol photons m^?2 s^?1, respectively). Long and short day lengths (16:8 and 8:16 LD cycles) were also applied at the two irradiances. Growth effects of temperature and irradiance were evident in most algae tested, and there were significant interactions among treatments. Most freshwater red algae had the best growth under low irradiance, confirming the preference of freshwater red algae for low light regimens. In general there was highest growth rate in long days and low irradiance. Growth optima in relation to temperature were species‐specific and also varied between low and high irradiances for the same alga. The most significant differences in pigment content were related to temperature, whereas few significant differences could be attributed to variation in irradiance and photoperiod or interactions among the three parameters. The responses were species‐specific and also differed for pigments in distinct temperatures, irradiances and photoperiods in the same alga. Phycocyanin was generally more concentrated than phycoerythrin and phycobiliproteins were more concentrated than chlorophyll a. The highest total pigment contents were found in two species typical of shaded habitats: A. hermannii and C. coeruleus. The expected inverse relationship of pigment with irradiance was observed only in C. coeruleus. In general, the most favorable conditions for growth were not coincident with those with highest pigment contents.     

Growth,toxicity and oxidative stress of a cultured cyanobacterium (Dolichospermum sp.) under different CO2/pH and temperature conditions

Cyanobacteria blooms are a worldwide nuisance in fresh, brackish and marine waters. Changing environmental conditions due to upwelling, changed mixing conditions or climate change are likely to influence cyanobacteria growth and toxicity. In this study, the response of the toxic cyanobacterium Dolichospermum sp. to lowered pH (?0.4 units by adding CO₂) and elevated temperature (+4°C) in an experimental set‐up was examined. Growth rate, microcystin concentration and oxidative stress were measured. The growth rate and intracellular toxin concentration increased significantly as a response to temperature. When Dolichospermum was exposed to the combination of elevated temperature and high CO₂/low pH, lipid peroxidation increased and antioxidant levels decreased. Microcystin concentrations were significantly correlated with growth rates. Our results show, although oxidative stress increases when exposed to a combination of high CO₂/low pH and high temperature, that growth and toxicity increase at high temperature, suggesting that the cyanobacterium in general seems to be fairly tolerant to changes in pH and temperature. Further progress in identifying biological responses and predicting climate change consequences in estuaries experiencing cyanobacteria blooms requires a better understanding of the interplay between stressors such as pH and temperature.     

野生稻近等基因系应答低温胁迫的生理生化指标分析

为了解水稻耐冷的生理机制,该文以野生稻近等基因系耐冷水稻品种DC907和其不耐冷受体亲本9311为材料,通过对水稻幼苗低温胁迫处理,检测分析了丙二醛(MDA)、超氧阴离子(O2-)、可溶性糖以及抗氧化酶类[超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)]等生理生化指标...     

Priming-induced changes in germination,morpho-physiological and leaf biochemical responses of fenugreek (Trigonella foenum-graecum) under salt stress

Abstract

Seed priming is a simple biotechnological tool which is potentially able to promote seed germination and invigoration as well as seedlings establishment and stress tolerance. In this study, the effects of seed pre-treatment with water (hydro-priming), 100 (HP-NaCl₁₀₀) and 200?mM (HP-NaCl₂₀₀) NaCl (halo-priming) for 6?h on some physiological and biochemical parameters of fenugreek was investigated under saline conditions (100?mM NaCl). For the three priming treatments, no significant changes in the final germination percentage were observed. However, a decrease in seed germination time was observed in hydro- and halo-primed (HP-NaCl₂₀₀) seeds. Salt stress (100?mM NaCl) reduced growth (shoot and root dry weight), pigment content, disturbed the ionic balance and enhanced malondialdehyde content. Salinity-induced changes in lipid metabolism towards synthesis/accumulation of saturated and monounsaturated fatty acids were observed in stressed plants. Seed priming ameliorated the negative effects of NaCl, ensuring significant amelioration on growth, pigment content, increased the activity of catalase and glutathione peroxidase and enhanced the synthesis of very long chain n-alkanes. Taken together, these data provide compelling evidence that priming is an effective alternative that can be used to promote germination and improves establishment and acclimation of fenugreek seedlings under saline conditions.     

Modelling temperature,photoperiod and vernalization responses of Brunonia australis (Goodeniaceae) and Calandrinia sp. (Portulacaceae) to predict flowering time

Background and Aims

Crop models for herbaceous ornamental species typically include functions for temperature and photoperiod responses, but very few incorporate vernalization, which is a requirement of many traditional crops. This study investigated the development of floriculture crop models, which describe temperature responses, plus photoperiod or vernalization requirements, using Australian native ephemerals Brunonia australis and Calandrinia sp.

Methods

A novel approach involved the use of a field crop modelling tool, DEVEL2. This optimization program estimates the parameters of selected functions within the development rate models using an iterative process that minimizes sum of squares residual between estimated and observed days for the phenological event. Parameter profiling and jack-knifing are included in DEVEL2 to remove bias from parameter estimates and introduce rigour into the parameter selection process.

Key Results

Development rate of B. australis from planting to first visible floral bud (VFB) was predicted using a multiplicative approach with a curvilinear function to describe temperature responses and a broken linear function to explain photoperiod responses. A similar model was used to describe the development rate of Calandrinia sp., except the photoperiod function was replaced with an exponential vernalization function, which explained a facultative cold requirement and included a coefficient for determining the vernalization ceiling temperature. Temperature was the main environmental factor influencing development rate for VFB to anthesis of both species and was predicted using a linear model.

Conclusions

The phenology models for B. australis and Calandrinia sp. described development rate from planting to VFB and from VFB to anthesis in response to temperature and photoperiod or vernalization and may assist modelling efforts of other herbaceous ornamental plants. In addition to crop management, the vernalization function could be used to identify plant communities most at risk from predicted increases in temperature due to global warming.     

Identification of a new‐to‐science cyanobacterium,Toxifilum mysidocida gen. nov. & sp. nov. (Cyanobacteria,Cyanophyceae)

Cyanobacteria occupy many niches within terrestrial, planktonic, and benthic habitats. The diversity of habitats colonized, similarity of morphology, and phenotypic plasticity all contribute to the difficulty of cyanobacterial identification. An unknown marine filamentous cyanobacterium was isolated from an aquatic animal rearing facility having mysid mortality events. The cyanobacterium originated from Corpus Christi Bay, TX. Filaments are rarely solitary, benthic mat forming, unbranched, and narrowing at the ends. Cells are 2.1 × 3.1 μm (width × length). Thylakoids are peripherally arranged on the outer third of the cell; cyanophycin granules and polyphosphate bodies are present. Molecular phylogenetic analysis in addition to morphology (transmission electron microscopy and scanning electron microscopy) and chemical composition all confirm it as a new genus and species we name Toxifilum mysidocida. At least one identified Leptolyngbya appears (based on genetic evidence and TEM) to belong to this new genus.     

Maize leaf temperature responses to drought: Thermal imaging and quantitative trait loci (QTL) mapping

Leaf temperature has been shown to vary when plants are subjected to water stress conditions. Recent advances in infrared thermography have increased the probability of recording drought tolerant responses more accurately. The aims of this study were to identify the effects of drought on leaf temperature using infrared thermography. Furthermore, the genomic regions responsible for the expression of leaf temperature variation in maize seedlings (Zea mays L.) were explored. The maize inbred lines Zong3 and 87-1 were evaluated using infrared thermography and exhibited notable differences in leaf temperature response to water stress. Correlation analysis indicated that leaf temperature response to water stress played an integral role in maize biomass accumulation. Additionally, a mapping population of 187 recombinant inbred lines (RILs) derived from a cross between Zong3 and 87-1 was constructed to identify quantitative trait loci (QTL) responsible for physiological traits associated with seedling water stress. Leaf temperature differences (LTD) and the drought tolerance index (DTI) of shoot fresh weight (SFW) and shoot dry weight (SDW) were the traits evaluated for QTL analysis in maize seedlings. A total of nine QTL were detected by composite interval mapping (CIM) for the three traits (LTD, RSFW and RSDW). Two co-locations responsible for both RSFW and RSDW were detected on chromosomes 1 and 2, respectively, which showed common signs with their trait correlations. Another co-location was detected on chromosome 9 between LTD and shoot biomass, which provided genetic evidence that leaf temperature affects biomass accumulation. Additionally, the utility of a thermography system for drought tolerance breeding in maize was discussed.     

高温胁迫对筛豆龟蝽成虫抗氧化能力的影响

为探索筛豆龟蝽Megacopta cribraria (Fabricius)对高温的耐受性及其体内抗氧化系统对极端高温胁迫的响应机制.以25℃为对照温度,经高温37℃、40℃、43℃和46℃胁迫处理4h后,测定筛豆龟蝽成虫存活率及其体内超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)的活性以及总抗氧化(T-AOC)能力和丙二醛(MDA)含量的变化.在37℃～46℃高温胁迫下,筛豆龟蝽成虫存活率与对照无显著差异,其体内SOD和CAT活性较对照组均显著增加,其中,SOD活性在37℃最高,随着温度进一步升高,其活性有所下降,但仍显著高于对照.CAT一直保持较高活性,随着处理温度的升高CAT活性呈逐渐升高趋势,在极端高温46℃时其活性最高.POD活性在37℃～ 40℃显著高于对照,43℃和46℃处理与对照无显著差异.T-AOC活性在各处理温度下均显著高于对照,且随着处理温度的升高T-AOC活性呈先升高后降低趋势.各处理MDA含量与对照无显著差异.筛豆龟蝽体内3种抗氧化酶的活力、总抗氧化能力和MDA在适应高温胁迫过程中发挥了重要作用.     

Phenology,irradiance and temperature characteristics of a freshwater red alga,Nemalionopsis tortuosa (Thoreales), from Kagoshima,southern Japan

Phenology, irradiance and temperature characteristics of a freshwater benthic red alga, Nemalionopsis tortuosa Yoneda et Yagi (Thoreales), were examined from Kagoshima Prefecture, southern Japan for the conservation of this endemic and endangered species. Field surveys confirmed that algae occurred in shaded habitats from winter to early summer, and disappeared during August through November. A net photosynthesis–irradiance (P–E) model revealed that net photosynthetic rate quickly increased and saturated at low irradiances, where the saturating irradiance (E_k) and compensation irradiance (E_c) were 10 (8–12, 95% credible interval (CRI)) and 8 (6–10, 95% CRI) μmol photon m^?2 s^?1, respectively. Gross photosynthesis and dark respiration was determined over a range of temperatures (8–36°C) by dissolved oxygen measurements, and revealed that the maximum gross photosynthetic rate was highest at 29.5 (27.4–32.0, 95%CRI) °C. Dark respiration also increased linearly when temperature increased from 8°C to 36°C, indicating that the increase in dark respiration at higher temperature most likely caused decreases in net photosynthesis. The maximum quantum yield (Fv/Fm) that was determined using a pulse amplitude modulated‐chlorophyll fluorometer (Imaging‐PAM) was estimated to be 0.51 (0.50–0.52, 95%CRI) and occurred at an optimal temperature of 21.7 (20.1–23.4, 95%CRI) °C. This species can be considered well‐adapted to the relatively low natural irradiance and temperature conditions of the shaded habitat examined in this study. Our findings can be applied to aid in the creation of a nature‐reserve to protect this species.