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Growth and photosynthetic activity of Sargassum henslowianum (Fucales, Phaeophyta) seedlings

Growth and photosynthetic activity of Sargassum henslowianum (Fucales, Phaeophyta) seedlings
Growth and photosynthetic activity of Sargassum henslowianum (Fucales, Phaeophyta) seedlings

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Growth and photosynthetic activity of Sargassum henslowianum (Fucales, Phaeophyta) seedlings in responses to different light intensities, temperatures and CO 2 levels under laboratory conditions

Binbin Chen a

& Dinghui Zou

ab

a

College of Environment and Energy , South China University of T echnology , Guangzhou,China

b

The Key Laboratory of Pollution Control and Ecosystem Restoration in Industry Clusters,Ministry of Education, Guangzhou, China Published online: 27 May 2014.

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ORIGINAL ARTICLE

Growth and photosynthetic activity of Sargassum henslowianum (Fucales,Phaeophyta)seedlings in responses to different light

intensities,temperatures and CO 2levels under laboratory conditions

BINBIN CHEN 1&DINGHUI ZOU 1,2*

1

College of Environment and Energy,South China University of Technology,Guangzhou,China,and 2The Key Laboratory of Pollution Control and Ecosystem Restoration in Industry Clusters,Ministry of Education,Guangzhou,China

Abstract

The habitat of Sargassum henslowianum (Fucales,Phaeophyta)is currently threatened by rising CO 2and global warming.Young seedlings of S.henslowianum regenerated from holdfasts were cultured under laboratory conditions with different levels of light intensity,temperature and CO 2concentration,aiming to examine the influences of these environmental factors on the growth and photosynthesis of this alga in its early growth stage.Young seedlings of Sargassum henslowianum exhibited faster growth under a light intensity of 60μmol photons·m ?2·s ?1than other light treatments (20and 120μmol photons·m ?2·s ?1).Seedlings growing at low light tended to develop blades and branches.Photosynthesis of seedlings increased with the temperature rising from 15to 25°C,and the blades or branches tended to develop with rising temperature.High temperature (30°C)depressed the growth of the young seedlings.Elevated CO 2significantly enhanced the germination vigor of the alga,but reduced net photosynthetic rate and irradiance saturation point.Our results suggest that experimental conditions,such as light intensities,temperature and CO 2levels each significantly affect the development and physiology of S.henslowianum seedlings.We propose that 60μmol photons·m ?2·s ?1and 25°C are preferable light –temperature conditions for the growth of S.henslowianum seedlings.

Key words :Climate change,environmental factors,photosynthesis,Sargassum henslowianum ,seedling

Introduction

The Sargassum genus,a group of brown algae without alternating phases in their life history,is the largest genus in the Phaeophyta and the most common brown alga in tropical and temperate regions around the world.The species of the Sargassum genus constitute an important part of the marine flora,forming extensive vegetation on shallow rocky coasts (Yoshida 1983;Guiry &Blunden 1991;Phillips 1995;Yoshida et al.1998;Haraguchi et al.2005).Some species of Sargassum exhibit strong tolerance and adaptive capacity to environmental factors and are endowed with the capacity to absorb various heavy metals (Kalyani et al.2004;Diniz &Volesky 2005;Rubin et al.2005).They contribute significantly to forming seaweed beds,balancing environment stress,and maintaining the stability and sustainability of ecosystems (Chopin 2001).

Recently,marine environments have been severely damaged by many kinds of human activity,especially development and utilization of coastal regions,which threaten the habitat of Sargassum species (Ang 2006).Additionally,atmospheric CO 2concen-tration has been rising continuously since the indus-trial revolution.Increased CO 2concentration in the atmosphere is causing global warming (Florides &Christodoulides 2009)and a decrease in the surface pH of oceans (Caldeira &Wickett 2003).Seawater acidification and increasing temperature caused by rising CO 2exerts strong effects on marine plants,such as the physiological status and chemical com-ponents of seaweeds (Zou &Gao 2010;Zou et al.2011b ).Meanwhile,light intensity vitally contributes

*Correspondence:Dr Dinghui Zou,College of Environment and Energy,South China University of Technology,Guangzhou,Guangdong 510006,China.E-mail:dhzou@https://www.wendangku.net/doc/f312147434.html,

Published in collaboration with the Institute of Marine Research,Norway Marine Biology Research,2014

Vol.10,No.10,1019–1026,

https://www.wendangku.net/doc/f312147434.html,/10.1080/17451000.2013.872798

(Accepted 15July 2013;Published online 26May 2014;Printed 2June 2014)?2014Taylor &Francis

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to seedling development and life history intergrada-tions of seaweeds and has considerable effects on the morphology and physiology of the marine macro-algae and the distribution of the intertidal seaweeds (Breeman &Ten Hoopen 1981;Davison &Pear-son 1996).

There have been some reports about the influence of various environmental factors on seedling growth of Sargassum algae,such as culture studies on the growth of four Sargassum species and Sargassum embryos in Hawaii (De Wreede 1976,1978)and ecological experiments with Sargassum muticum (Yendo)Fensholt in the Pacific (Norton 1977;Hales &Fletcher 1989).Steen (2004)reported the effects of reduced salinity on reproduction and germling development in S.muticum .Ang (2006)reported the phenology of Sargassum spp.in Tung Ping Chau Marine Park in Hong Kong.Hwang et al.(2006)also researched artificial seed production and cul-tivation of the edible Sargassum fulvellum (Turner)C.Agardh in Korea.Wong &Phang (2004)studied the seasonality in biomass production of Sargassum baccularia (Mertens) C.Agardh and Sargassum aquifolium (Turner)C.Agardh at Cape Rachado in Malaysia and found that rainfall was the most important factor that controlled the biomass pro-duction and reproduction for both species.

Studies on the reproductive biology of Sargassum spp.,especially on their early sexual propagation and germling development,are relevant to basic and applied research in the Sargassum genus.Studies on early growth stages of Sargassum seedlings and seedling technology have been carried out as stated above,but these studies were limited to a few species.Moreover,the classification,distribution and population status of many species are currently not completely clear.The species Sargassum henslo-wianum C.Agardh (Fucales,Phaeophyta)has extre-mely important ecological and economic value.Zou et al.(2011a )compared the photosynthetic carbon acquisition of S.henslowianum between the vegetat-ive and reproductive tissues.Wei et al.(2012)studied the chemical constituents of the brown alga S.henslowianum collected from the South China Sea.However,to date,there has been no research on the influence of environmental factors on early S.henslowianum seedling development.We collected wild S.henslowianum and obtained seedlings from the plant thalli in the holdfasts.Then culture experiments of the seedlings with different light intensity,temperature and CO 2levels were carried out under laboratory conditions,aiming to examine the influence of important environmental factors on the development and physiology of S.henslowianum at an early growth stage.

Material and methods

Seaweed material and experimental treatments

Sargassum henslowianum was collected from lower intertidal rocks on the coast of Nan ’ao Island,Shantou,China (23°20′N,116°55′E).Visible epi-phytes and attached sediments were cleaned from thalli with natural seawater,and the samples were transported to the laboratory in an insulated polysty-rene cooler (about 5°C).Healthy algae were selected and rinsed in sterile seawater and were maintained in a glass aquarium containing natural,filtered seawater with extra nitrogen and phosphorous (salinity ~33psu,~20μmol·l ?1NO 3?,~1μmol·l ?1PO 43?)in CO 2incubators (GXZ-300D,Jiangnan Instrument Fact-ory,Ningbo,China).Culture conditions were main-tained at 20°C and 30–40μmol photons·m ?2·s ?1(fluorescent illumination)with 14:10h (light:dark)photoperiod to induce the maturation of the holdfasts and regeneration of seedlings.The seedlings with about 2mm main axis were collected after 20dayas.Then they were cultured in CO 2incubators with continuous aeration.Basic culture conditions were maintained as follows:a sample of 0.2g fresh weight was cultured at 20°C in 1l filtered natural seawater with extra nitrogen and phosphorous (salinity ~33psu,20μmol·l ?1NO 3?,1μmol·l ?1PO 43?)in glass culture tanks.The illumination was controlled at ~80μmol photons·m ?2·s ?1by fluorescent tubes for 12:12h (light:dark)photoperiod.The culturing con-ditions were changed according to experiment proto-col,and culture media were renewed every 2days.Experiment I:effect of light intensity

The illumination experiment was carried out by controlling the intensity of fluorescent tubes while culturing the seedlings.The fluorescent illu-mination intensities were maintained at 20μmol photons·m ?2·s ?1(low light,LL),60μmol photons·m ?2·s ?1(intermediate light,IL)and 120μmol photons·m ?2·s ?1(high light,HL)at 20°C.The other cultivation conditions were maintained as basic culture conditions as described above.The physiological traits (growth and photosynthesis)of the samples were determined after 10days of laboratory culture.

Experiment II:effect of temperature

In the temperature experiment,temperatures in the CO 2incubators were maintained at 15,20,25and 30°C at ~80μmol photons·m ?2·s ?1light intensity,respectively.The other culture conditions were main-tained as basic culture conditions.The physiological traits of each sample were determined after 10days of culture.

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Experiment III:effect of CO 2level

The CO 2concentrations were controlled at 390μl·l ?1(ambient air,AA)and 750μl·l ?1(elevated air,EA)in two CO 2incubators,the illumination was maintained at ~80μmol photons·m ?2·s ?1by fluorescent tubes at 20°C,and culture medium was renewed every 2days.Other cultivation conditions were maintained as basic culture conditions.The physiological traits of each sample were determined after 10days of indoor culture.

Relative growth rate

Changes in biomass (fresh weight,F W )were meas-ured at the end of the culture period to estimate growth.Mean relative growth rate (RGR )was calculated using the formula:RGR (%day ?1)=ln (W t /W 0)/t ×100,where W 0referred to the initial F W ,and W t referred to the F W after t days.

Germination vigour

Germination vigour was originally driven by the ability of the plant embryo,embedded within the seed,to resume its metabolic activity in a coordi-nated and sequential manner (Rajjou et al.2012).Sixteen Sargassum henslowianum seedlings were ran-domly selected during the initial and final stages of each treatment,and the blades (or branches)and sprouts of each were counted (G ).Germination vigour (Gv )was calculated using the formula:Gv (%day ?1)=ln(G t /G 0)/t ×100;in this formula,G 0referred to initial G ,and G t referred to the G after t days.

Photosynthetic oxygen evolution and respiration rate The net photosynthetic O 2evolution rate (NPR )of Sargassum henslowianum seedlings at different photon irradiances (P vs.I curves)was determined by using a Clark-type oxygen electrode (YSI Model5300,Yellow Springs Instrument Co.,OH,USA),with a water jacket connected to a cooling circulator for maintaining the temperature at the desired level.Illumination was provided by a hal-ogen lamp.Irradiance,which was measured with a quantum sensor (SKP 200,ELE International,Leighton Buzzard,UK)at the surface of the cham-ber was,respectively,set at eight levels from 0to 800μmol·m ?2s ?1photon fluence by altering the distance between the electrode chamber and the light source.The electrode chamber contained 8ml original culture solution and was magnetically stirred.Dark respiration rate (R d )was obtained by determining dark O 2consumption.Approximately 0.06g (fresh weight,F W )of sample was utilized in

each test.NPR and dark respiration were expressed as μmol O 2·g ?1Fw·h ?1.Photosynthetic parameters were calculated based on Henley (1993).

Statistical analyses

Origin 8.0(Origin Lab Corp.,Northampton,MA,USA)was used in data processing and statistical analysis.One-way analysis of variance and the Tukey test were used to analyse differences among treat-ments.Tests for normality and homogeneity of variance were performed to check assumptions of parametric analysis.All the data were expressed as means ±SD (n ≥3).A P -value of 0.05was considered to be statistically significant.

Results

Experiment I

Light intensity exerted a significant influence on the development of Sargassum henslowianum seedlings in the early growth stage during the laboratory incuba-tion (Figure 1a ).Relative growth rate (RGR )of the seedlings cultured under intermediate light (60μmol photons·m ?2·s ?1)was considerably higher (F 2,6=40.23,P <0.01)than that of other treatments (i.e.treatments of 20and 120μmol photons·m ?2·s ?1).The seedlings growing at low light (20μmol photons·m ?2·s ?1)exhibited the highest value of germination vigour (Gv )(F 2,6=50.97,P <0.01)among the three light intensity treatments.RGR and Gv of the seedlings cultured under high light (120μmol photons·m ?2·s ?1)were evidently lower than those under intermediate light and low light.

Figure 1b showed the net photosynthetic O 2evolution rates (NPR )at different photon irradiances (P –I curves)for young S.henslowianum seedlings grown at different light intensities,and the photo-synthetic parameters of P –I curves are presented in Table I .For all the light treatments,no photoinhibi-tion was observed at the maximum illumination intensity of 800μmol photons·m ?2·s ?1.Among all the light intensity treatments,S.henslowianum seed-lings growing at intermediate light intensity (60μmol photons·m ?2·s ?1)exhibited the highest irradiance-saturated net photosynthetic rate (NPR m ;F 2,6=14.34,P <0.01).There were no statistically signi-ficant differences in respiration (R d ;F 2,6=3.56,P =0.10),irradiance saturation point (I k ;F 2,6=1.02,P =0.42),apparent photosynthetic efficiency (α;F 2,6=4.76,P =0.06)or compensation irradiance point (I c ;F 2,6=0.86,P =0.47)among the seedlings growing at different light levels.

Growth and photosynthesis of Sargassum henslowianum seedlings

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Experiment II

Temperature had a significant effect on the growth of young Sargassum henslowianum seedlings (Figure 2a ).RGR of seedlings significantly increased as the tem-perature rose from 15to 25°C (F 3,8=39.35,P <0.01),but it was drastically reduced at 30°C (F 1,4=187.00,P <0.01).The Gv of seedlings was similar

between 15and 25°C (F 3,8=35.78,P <0.01),whereas it was lowered at high temperature (30°C;F 1,4=96.31,P <0.01).

The NPR m value significantly increased as the temperature rose from 15to 25°C (F 2,6=59.02,P <0.01;Table II ,Figure 2b ).At 30°C,NPR m was considerably reduced compared with that of the algae growing at 25°C (F 1,4=184.79,P <0.01).The respiration (F 2,6=9.25,P <0.05),α(F 2,9=26.75,P <0.01)and I k (F 2,6=20.09,P <0.01)increased with rising temperature,and these parameters reached a maximum at 25°C.There was no signific-ant difference of I c among all the temperature treat-ments (F 3,8=2.03,P =0.19).

Experiment III

Elevated CO 2(EA)did not significantly enhance the growth of young Sargassum henslowianum seedlings (F 1,4= 3.18,P =0.15),as shown in Figure 3a .However,the Gv of seedlings cultured under ele-vated CO 2was significantly greater than that under ambient air (AA;F 1,4=23.79,P <0.01).

The young seedlings growing at ambient air exhib-ited a higher NPR m value than the thalli growing

at

Figure 1.Relative growth rates (RGR )and germination vigour (Gv )(a)and rates of photosynthetic O 2evolution vs.irradiance (P –I)curves (b)of young Sargassum henslowianum seedlings cultured under different light levels (20μmol photons·m ?2·s ?1,LL;60μmol photons·m ?2·s ?1,IL;120μmol photons·m ?2·s ?1,HL).Vertical bars represent ±standard deviation (SD)of the means (RGR :n =3,Gv :n =3,NPR :n =

3).

Figure 2.Relative growth rates (RGR )and germination vigour (Gv )(a)and rates of photosynthetic O 2evolution vs.irradiance (P –I)curves (b)of young Sargassum henslowianum seedlings cultured under different temperatures (15,20,25,30°C).Vertical bars represent ±standard deviation (SD)of the means (RGR :n =3,Gv :n =3,NPR :n =3).

Table I.Effects of different light intensities (20μmol photons·m ?2·s ?1,LL;60μmol photons·m ?2·s ?1,IL;120μmol photons·m ?2·s ?1,HL)on photosynthetic parameters of P –I curves of young seedlings of Sargassum henslowianum .

LL

IL HL NPR m 23.01±4.0934.05±5.7215.25±2.57R d ?3.20±1.20?4.14±1.34?1.70±0.76α0.15±0.050.19±0.010.12±0.02I c 21.74±7.9121.05±6.2315.31±5.35I k

138.02

±37.42

152.48

±17.78

124.43

±5.25

NPR m ,Maximum net photosynthetic rate (μmol O 2·g ?1F W ·h ?1);R d ,dark respiration rate (μmol O 2·g ?1F W ·h ?1);α,apparent photosyn-thetic efficiency [(μmol O 2·g ?1F W ·h ?1)/(μmol photons·m ?2·s ?1)];I c ,compensation irradiance point (μmol photons·m ?2·s ?1)and I k ,irradiance saturation point (μmol photons·m ?2·s ?1).Values are means ±SD (n =3~4).

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elevated CO 2(F 1,4=11.03,P <0.05;Figure 3b ).Elevated CO 2reduced the irradiance saturation point (I k )of young S.henslowianum seedlings (F 1,4=21.22,P <0.01),but hardly affected the respiration (R d ;F 1,4=0.12,P =0.75),apparent photosynthetic efficiency (α;F 1,6=0.05,P =0.83)or compensation irradiance point (I c ;F 1,4=0.18,P =0.70;Table III ).Discussion

Influence of light intensity

The results showed that the growth of Sargassum henslowianum seedlings in early growth stages was accelerated when the light intensity rose from 20to 60μmol photons·m ?2·s ?1.Seedlings growing at low light levels tended to develop blades and branches,while high light levels depressed the growth of S.henslowianum seedlings.Intermediate light enhanced photosynthesis and respiration of S.henslowianum seedlings,while the high light level lowered the maximum net photosynthetic rate,apparent photosyn-thetic efficiency,compensation irradiance point,and irradiance saturation point of the thalli.

Young seedlings of S.henslowianum cultured under 60μmol photons·m ?2·s ?1exhibited a greater NPR and RGR than other light intensity treatments,and reached the maximum value of NPR m .It was reported that the optimal irradiance intensity for growth of Sargassum thunbergii (Mertens ex Roth)Kuntze germlings (Zhao et al.2008)and S.muticum at 25°C and 34‰salinity (Hales &Fletcher 1989)was 44μmol photons·m ?2·s ?1.The growth of Sargassum species may take advantage of the relative low light intensity.An illumination intensity of 60μmol photons·m ?2·s ?1was considered a preferable light intensity for the growth of S.henslowianum seedlings in the early growth stage.Hales &Fletcher (1989)reported that the germlings of S.muticum had a broad tolerance to irradiance from 9to 175μmol photons·m ?2·s ?1.We also found a broad light tolerance in S.henslowianum seedlings,although there was a clear optimum for growth at 60μmol photons·m ?2·s ?1.The light intens-ities of 20and 120μmol photons·m ?2·s ?1,especially a

light intensity of 120μmol photons·m ?2·s ?1,were considered as thresholds for growth and photosynthesis of S.henslowianum in early growth stage,and the light intensity of 60μmol photons·m ?2·s ?1was an optimal and preferable light intensity for S.henslowianum seed-ling culture.

Influence of temperature

When the light intensity was maintained at 80μmol photons·m ?2·s ?1,with a raised temperature (from 15to 25°C),the growth of young S.henslowianum

Table II.Effects of different temperatures (15,20,25,30°C)on photosynthetic parameters of P –I curves of young seedlings of Sargassum henslowianum .

15°C

20°C 25°C 30°C NPR m 14.92±3.9734.05±5.7255.18±3.6617.67±3.08R d ?3.79±1.70?4.14±1.34?8.44±1.36?3.05±0.96α0.10±0.020.19±0.010.23±0.040.12±0.01I c 35.80±11.8621.05±6.2334.26±1.9829.97±8.71I k

107.35

±8.37

152.48

±17.78

191.71

±20.30

121.56

±25.68

NPR m ,Maximum net photosynthetic rate (μmol O 2·g ?1F W h ?1);R d ,dark respiration rate (μmol O 2·g ?1F W h ?1);α,apparent photosynthetic efficiency [(μmol O 2·g ?1F W h ?1)/(μmol photons·m ?2·s ?1)];I c ,compensation irradiance point (μmol photons·m ?2·s ?1)and I k ,irradiance saturation point (μmol photons·m ?2·s ?1).Values are means ±SD (n =3~

4).

Figure 3.Relative growth rates (RGR )and germination vigour (Gv )(a)and rates of photosynthetic O 2evolution vs.irradiance (P –I)curves (b)of young Sargassum henslowianum seedlings cultured under ambient air (390μl·l ?1)and elevated CO 2(750μl·l ?1).Vertical bars represent ±standard deviation (SD)of the means (RGR :n =3,Gv :n =3,NPR :n =3).

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seedlings was enhanced.Many Sargassum species have a wide temperature growth range.The length of S.thunbergii germlings grown at 88μmol photons·m ?2·s ?1increased as the temperature rose from 10to 25°C (Zhao et al.2008).The filamentous gametophytes of the algae can grow at temperatures of 8–28°C and can recover from conditions of ?1to 33.5°C (Sanderson 1990;Peters &Breeman 1992).Hales &Fletcher (1989)also found a positive relationship between growth of S.muticum germ ‐lings and a temperature increase from 10to 20°C.

Low temperature depressed the growth and ger-mination vigour of S.henslowianum seedlings.Low temperature (10°C)also reduced germling growth of S.muticum as reported by Zhao et al.(2008).However,a high temperature (30°C)hardly enhanced the growth and germination of the S.henslowianum seedlings.A temperature of 25°C was preferable for S.henslowianum growth in early stages.Distinct critical temperatures also exist in other Sargassum species.Temperatures above 26°C were reported to hamper the development of Sargassum fusiforme Harvey (Setchell)in the young germling stage (Sun et al.1996;Hwang et al.1997).A temperature of 24°C was considered to be near the optimum for the development of Sargassum spp.embryos (De Wreede 1976).Haraguchi et al.(2005)found that the maximum critical temperatures were 27°C for Sargassum horneri (Turner)C.Agardh,S.hemiphyllum (Turner) C.Agardh and S.micra-canthum (Kützing)Endlicher,30°C for S.piluliferum (Turner) C.Agardh,S.fulvellum (Turner) C.Agardh and Myagropsis myagroides (Mertens ex Turner)Fensholt and 31°C for S.patens C.Agardh,S.macrocarpum C.Agardh and S.thunbergii .

There are few reports in the literature on the influence of temperature on the photosynthesis of Sargassum germlings.It has been reported that the photosynthetic rates of Undaria pinnatifida (Harvey)

Suringar gametophytes,which also belong to the Phaeophyta,declined but remained positive at incubation temperatures from 17to 31°C by utiliz-ing heat shock proteins (Hsps)via a mechanism that was downstream of gene transcription,while respira-tion increased with increasing temperatures up to 26°C and then declined (Henkel &Hofmann 2008).The oxygen consumption of the kelp Macrocystis pyrifera (Linnaeus)C.Agardh gametophyte dropped from approximately 25to ?25mg O 2·g ?1·F W ·h ?1when temperature increased from 15to 30°C (Fain &Murray 1982).The photosynthesis and respiration of S.henslowianum germlings were enhanced as the temperature rose from 15to 20°C.High temperat-ure (30°C)depressed the photosynthesis and res-piration of the germlings ’thalli.In this research,young germlings of the macroalga S.henslowianum have been shown to have a broad temperature range for photosynthesis and survival,as in other Sargas-sum species (Hales &Fletcher 1989;Hwang et al.2006;Zhao et al.2008).

The results of our temperature experiments indi-cated that seedlings of S.henslowianum in their early growth stage showed temperature tolerance of 15–25°C,which is similar to that of S.thunbergii (Haraguchi et al.2005).They exhibited the ability to withstand temperatures that might be experienced in the future due to global warming (20–25°C).The upper temperature threshold for S.henslowianum seedlings is likely to be around 30°C,while the optimum temperature range for culture growth is 20–25°C at 80μmol photons·m ?2·s ?1.

In the present study,severe coalescence (adhesion)was observed in our low temperature culture in the temperature experiments.Early stages of sporeling coalescence was observed in Gracilaria chilensis C.J.Bird,McLachlan & E.C.Oliveira and polysporic plantlets were formed by groups of spores and their derivatives (Santelices et al.1996).The frequency of spore coalescence in field populations of species of Gracilaria and other gregarious Rhodophyta species would further increase the possibilities for individual and population survival,either by increased variation and resistance to the abiotic environment,by increased competitive capacity or by increased resistance to grazers (Buss 1985;Maggs &Cheney 1990;Paine 1990);therefore,coalescence was suggested as being an ecologically important adaption of clonal growth (Santelices et al.1996).In this research,the coales-cence phenomenon of young seedlings of S.henslowia-num was probably due to the low temperature,which depressed the rapid elongation of the thalli;conse-quently,the growing rhizoid coalesced the seedlings which adhered to each other.As the temperature rose,the blade or branch of the seedlings tended to develop.

Table III.Effects of different CO 2levels in culture (ambient air,390μl·l ?1;and elevated CO 2,750μl·l ?1)on photosynthetic parameters of P –I curves of young seedlings of Sargassum henslowianum.

390μl·l ?1

750μl·l ?1NPR m 34.05±5.7221.34±3.35R d ?4.14±1.34?4.50±1.19α0.19±0.010.19±0.02I c 21.05±6.2323.50±8.02I k

152.48

±17.78

87.23

±16.91

NPR m ,Maximum net photosynthetic rate (μmol O 2·g ?1F W h ?1);R d ,dark respiration rate (μmol O 2·g ?1F W h ?1);α,apparent photosyn-thetic efficiency [(μmol O 2·g ?1F W h ?1)/(μmol photons·m ?2·s ?1)];I c ,compensation irradiance point (μmol photons·m ?2·s ?1)and I k ,irradiance saturation point (μmol photons·m ?2·s ?1).Values are means ±SD (n =3~4).

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Influence of CO 2level

Elevated CO 2significantly enhanced the blade ger-mination of Sargassum henslowianum seedlings,but reduced the photosynthesis and irradiance saturation point of the thalli.In contrast,light-saturated net photosynthesis increased with rising CO 2concentra-tions and became saturated at CO 2concentrations higher than the present atmospheric level in S.hemiphyllum during emersion,as reported by Zou &Gao (2005).Oxygen evolution rates at pH 8.1were nearly saturated with the 2.2mM inorganic carbon (Ci;the normal concentration in seawater)for both the blade and receptacle of S.henslowianum ,but increased in an almost linear way with the increasing Ci concentrations over the range from 1.1to 15.4mM at pH 9.0and with lowered pH levels from 8.0to 6.0the NPR of the alga for both the blade and receptacle tissues was gradually depressed (Zou et al.2011a ).This was possibly due to the presence of an active CCM (carbon concentrating mechanism),which is mainly related to efficient HCO 3?utilization (Zou et al.2011b ).Moreover,increased CO 2concentration resulted in the pH value decreasing (Caldeira &Wickett 2003).An alteration of pH created a major influence on the macroalgal growth and Ci utilization (Mercado &Niell 1999;Zou et al.2011a ).In our research,reduction of the photosynthesis and I k of the young seedlings probably resulted from the decreased pH in seawater,induced by elevated CO 2,and the lowered pH depressed the photosynthesis and res-piration of the thalli.The increased development of the blade germination of S.henslowianum seedlings was probably due to the elevated HCO 3?resulting from elevated CO 2.Conclusions

We conclude that enhancement of intermediate light intensity (60μmol photons·m ?2·s ?1)on the growth of holdfast-derived young seedlings of Sargassum henslowianum was greater than that of other light treatments;seedlings growing at low light tended to develop blades or branches.The optimum temper-ature for photosynthesis and growth of S.henslowia-num seedlings was 25°C.Elevated CO 2(750μl·l ?1)significantly enhanced the germination vigour of the seedlings,but reduced the photosynthesis and irradi-ance saturation point of the young thalli.Our results suggested that experimental conditions,such as light,temperature and CO 2levels,significantly affected the development and physiology of S.henslowianum in the early growth stage,but the interaction of these environmental factors should be further investigated.Based on our study,we suggest

the optimum culture conditions for S.henslowianum seedlings are 60μmol photons·m ?2·s ?1and 25°C.

Funding

This study was supported by the National Natural Science Foundation of China (41276148and 41076094).

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Growth and photosynthesis of Sargassum henslowianum seedlings

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Editorial responsibility:Mirta Teichberg

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精品—忠言逆耳助我行作文-忠言逆耳助我行议论文三篇

精品—忠言逆耳助我行作文|忠言逆耳助我行议论文三篇 忠言逆耳助我行议论文(一) 在我小时候,因为不听妈妈的话,结果差点就丢了小命,到至今都有些后怕。 事情是这样的,那是一年夏天的中午,我和姐姐吃晚饭后,就在一起嘀嘀咕咕的商量着一会到河边去玩会水,我和姐姐正说着,姐姐正说着,妈妈突然拍了拍我们说:热了在家洗洗澡,我和姐姐拿起书包就跑,我们两个一看,氺好深!我和姐姐不约而同的喊了一声,然后扔下书包就往水边跑,然后各自坐在了一边,脚垂到水里,在水中脚溅起的浪花,在加上又刮来一清风,感觉凉快多了。过了一会姐姐说:你往那边去去,我往你那边坐坐。我猛地点点头,然后小心地往那边移了移,谁知,突然一只脚踩空了,而后陷进了泥土里不知不觉,另一只脚就漂了起来,然后我才知道我掉进了水里,就要刮跑了。这时我条件反射似的一只手突然从水里伸了出来,当时的水都把我整个人都埋住了,姐姐看到这个情况,吓了一跳,赶紧拉着我的手,特别有力的把我往上拉,嘴里还喊着上来,快上来。这时姐姐班的几个男生从旁边经过,开玩笑的说了生:哎,你在捞鱼啊!我妹掉里面了,快过来帮帮忙。听到姐姐说有人掉河里了,那个男生立马跑过来,叫我:哎,把你另一只手伸出来。当时我听到了那个声音,就赶快把手伸了出去,在姐姐和那位哥哥地帮助下,我被拉了出来,看着那位哥哥,我个姐姐说:谢谢你!而那位哥哥却说:没事,以后不要再来这里玩了。然后就走了。但是我的鞋子也被刮跑了一只,然后又跑回家换了一双。不过谁知这件事让妈妈知道了,然后狠狠地教训了我们一下,生气地说:跟你们说了多少遍了,不准往河边去,就在家洗洗,不听话,这次知道了吧!我和姐姐都乖乖地点了点头。从那以后我和姐姐就听了妈妈的话,再也没有去河边玩了。然后我也受到了教训,现在只要有深水,我就特别地害怕。 经过了这件事,我记得了妈妈地许多不可以,因为那些不可以,都是对我好,那些逆耳忠言,也是对我好,它们就像一盏盏指路的明灯,指引我们走向正确的道路。 忠言逆耳助我行议论文(二) 曾几何时,在流言蜚语中派回,找不到方向;曾几何时,在他人的言语中迷茫,不知路在何方。只因为没有忠诚的劝诫,前途一片迷茫。直到现在,才懂得逆耳忠言对我们的谆谆教育。 不曾忘却:忠言逆耳利于行,良药苦口利于病。回眸史册,商纣王,不听忠诚劝告,亲手把自己的国家埋葬;蔡恒公不听扁鹊的建议,最后只能把自己的命搭上。 有人说,那已成为历史,距现在已经相当遥远,不知道的参考。那么说说在我身上所发生的吧! 上中学后,我的几个好朋友都迷上了网络游戏,经常三五成群偷偷的跑去上网,他们也常常告诉我网络游戏如何好玩,在里面想干什么就干什么无不痛快。终于,在他们的勾引下,我也步入其中。刚开始,只觉得好玩,便三番两次的和他们同去,渐渐地我也迷恋上了她。 可是好运总不能永远陪伴我。有一次,我和他们玩得正高兴的时候,老师来了。老师对我说:千万不要网络游戏迷惑了你们,那样,他会毁了你们的一生。我明白老师是为我们好的,可是一旦迷恋上她,就别想一下摆脱它。我仍然是我行我素,只是去的次数逐渐减少。 直到发生了一件事,我才彻底摆脱它。那是星期六的一天,我有去玩了。却没想到爸爸也跟着我,我刚到门口,爸爸的大手拉着我往家去。回到家中,免不了皮肉之苦,过了一会儿,爸爸的气消了,对我一阵心灵教育:孩子,你知道吗,网络游戏对你来说有什么好处呢,网络是一把双刃剑,你总是看到它坏的一面。你自己好好想想,你今后怎么办吧。 从那以后,我再也没有越雷池半步。而同伴的成绩一次不如一次。我这才惊觉老师,爸爸的忠言在帮助我们抵制不良诱惑啊。 忠言逆耳利于行。我终于明白了这个道理。 忠言逆耳助我行议论文(三)

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古代汉语词类活用例句列举 古代汉语词类活用例句列举《郑伯克段于鄢》1、例:壮公生,惊姜氏。P97 惊:用作使动,使。。。惊。2、例:无生民心。P99 生:用作使动,使。。。产生。3、例:若阙地及泉,隧而相见。P101 隧:名词动用。《公孙无知之乱》4、豕立而啼,P109 立:名词作状语,像人一样丫立。〈安之战〉5、皆主?献子。P117 主:名词动用,以。。。为主。6、君无所辱命。P119 辱:动词使动,使。。。受辱。7、从左右,皆肘之。P123 肘:名词使动,表示用胳膊推撞。8、臣辱戎士。123 辱:动词使动。9、人不难以死免其君。P123 免:用作使动,使。。。免于。10、故中御而从齐候。P123 中:方位名词做状语。〈子产说范宣子轻敝〉11、三周华不注。P122 周:

名词动用。12、郑人病之。P129 病:名词用作意动。13、象有齿而焚其身。P130 焚:动词用作使动。14、宣子说,乃轻弊。P130 轻:形容词用作使动,使。。。轻。〈苏秦连横约纵〉15、今先生俨然不运千里而庭教之。P182 远:形容词用作意动。16、明言章理,兵甲愈起。P183 明、章:用作使动。 1 17、辨言伟服。攻战不息。P183 辩、伟:都用作使动,使。。。雄辩,使。。。华美。18、繁称文辞,天下不冶。P183 文:名词用作使动。19、夫徒处而致利,安坐而广地。P183 广:形容词用作使动,使。。。广。20、言语相结,天下为一。P183 言语:名词作状语。21、今欲并天下,凌万乘,诎敌国。制海内,子元元。臣诸候。非兵不可。P183 诎:用作使动,使。。。屈服;子:名词用作使动,使。。。成为子女;臣:名词用作使动,使。。。成为臣子。22、约纵散横,以抑强秦。

忠言逆耳利于行精选作文

忠言逆耳利于行精选作文 篇一:忠言逆耳利于行 我们每个人都爱听好听话,但是人生道路不可能事事都一帆风顺。假如我们一直伴着好听话成长,一直执迷不悟地沉醉于自己所谓的一点点成绩,那么我们就永远不可能成长。成长的道路上,一定会有人在我们的耳边说着不好听的话,说着令我们难堪的话,或长辈,或兄弟姐妹,或朋友。我们也许会生气,也许会厌恶,但是这些不那么好听的话却能激励我们进步,教诲我们成长,这就是——忠言逆耳利于行。 唐代的名臣魏征经常向唐太宗进谏,甚至有时当面指出唐太宗的不足,弄得唐太宗经常下不来台,甚至扬言要杀掉魏征。但是就是这样的忠臣,这样逆耳的话语,却时时刻刻提醒着唐太宗,勉励着唐太宗。魏征的《谏太宗十思疏》,唐太宗每天早上朗读它一遍,因为它无时无刻不催促着唐太宗为民造福,做一个明君。这是忠言逆耳利于行的正面例子。反之,历史上也一定有反面例子。比如比干劝商纣王不能横征暴敛反被剖心,伍子胥劝吴王杀掉勾践夫妇不成反被杀害。所以忠言不是对所有人都能产生积极影响,有的人一点就通,能够虚心接受别人的意见,有的人却夜郎自大,至死不悟。 但是现代社会和古代不同,我们更应该认真聆听逆耳忠言,更加

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前的我了,上课我会很专心地听讲,吸收老师所传授的知识,回家后也暗暗努力地学习。终于我等到了机会,在期中考,我取得了惊人的进步。可我却没有表现出太多的惊喜,我继续努力着,努力着……就在我一次次得到优异的成绩后,我收敛了我的叛逆,我只在老师的办公桌上放了一张字条“我只想证明给某些人看,他的看法是错误的!”成功的喜悦冲昏了我的头脑,以至于我没有留意到老师对我欣慰的嘴角上扬的弧度…… 自从小学毕业后,我就再也没有回去母校,因为那里有太多不好的回忆。可我却不知道,由于那段奋斗的时光让我能够比别人更快地适应中学繁忙的学习生活,而这一切,都是我一直所“讨厌”的老师给予的。 “良药苦口利于病,忠言逆耳利于行,妈,我知道了。”我将手中的药一饮而尽,然后向母校的方向跑去……

初中所学文言文中的五类常见词类活用现象

初中所学文言文中的五类常见词类活用现象

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①代词前面的名词用如动词(肘之、面之),因为代词不受名词修饰; ②副词尤其是否定副词后面的名词用如动词(“遂东”、“不君”); ③能愿动词后面的名词也用如动词(“能水”、“欲刃”); ④句中所确定的宾语前面的名词用如动词(“脯鄂侯”“手剑”) (二)动词、形容词、名词的使动用法 一、动词的使动用法。 定义:主语所代表的人物并不施行这个动词所表示的动作,而是使宾语所代表的人或事物施行这个动作。例如:《左传隐公元年》:“庄公寤生,惊姜氏。”这不是说庄公本人吃惊,而是说庄公使姜氏吃惊。 在古代汉语里,不及物动词常常有使动用法。不及物动词本来不带宾语,当它带有宾语时,则一定作为使动用法在使用。如: 焉用亡.郑以陪邻?《左传僖公三十年》 晋人归.楚公子榖臣与连尹襄老之尸于楚,以求知罃。(左传成公三年) 大车无輗,小车无杌,其何以行.之哉?《论语·为政》 小子鸣.鼓而攻之可也。《论语·先进》 求也退,故进.之;由也兼人,故退.之。《论语·先进》 故远人不服,则修文德以来.之。《论语·季氏》 有时候不及物动词的后面虽然不带宾语,但是从上下文的意思看,仍是使动用法。例如《论语·季氏》:“远人不服而不能来也”这个“来”字是使远人来的意思。 古代汉语及物动词用如使动的情况比较少见。及物动词本来带有宾语,在形式上和使动用法没有什么区别,区别只在意义上。使动的宾语不是动作的接受者,而是主语所代表的人物使它具有这种动作。例如《孟子·梁惠王上》“朝秦楚”,不食齐宣王朝见秦楚之君,相反的,是齐宣王是秦楚之君朝见自己。 下面各句中的及物动词是使动用法: 问其病,曰:“不食三日矣。”食.之。《左传·宣公二年》

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太叔却以宽政治民,致使盗贼横行;夫差不听信伍子胥之忠言,终被勾践灭国。倘若他们听取了忠告,三国也许会提前结束,子太叔也许会与子产齐名,夫差也就不会亡国。因为忠言与他们内心的想法不符,就回绝其言,往往会引人走上失败之路。由此可见,忠言虽然逆耳,却能打开成功之门,避开失败的深渊。 当我们走在追求成功的路上的时候,请谨慎对待那些逆耳的建议,因为只有它们,才能给予我们真正的成功之路,当你拒绝了它们,也就亲手关闭了成功的大门,所以请接受逆耳忠言。

词类活用例子

文言实词词类活用 活用为一般动词 (一)名词活用为一般动词 1.两个名词连用,既不是并列关系,又不是修饰关系,便是动宾或主谓,其中一个必然活用为动词。 a .有一老父,衣褐,至良所。 b.籍吏民,封府库。 c.我有嘉宾,鼓瑟吹笙。 d.冬雷震震夏雨雪。 2.名词后紧跟代词,该名词活用为动词。 a.驴不胜怒,蹄之。 b.以其乃华山之阳名之。 c.名余曰正则兮。 3.名词放在副词后,便活用为动词。 a.日将暮,取儿槁葬。 b.太子及宾客知其事者,皆白衣冠以送之。 c.从弟子女十人所,皆衣缯单衣,立大巫后。 4.名词放在“能”“可”“足”“欲”等呢过愿动词后,便活用为动词。 a.假舟楫者,非能水也。 b.云青青兮欲雨。 c.其力尚足以入,火尚足以明。 d.子谓公冶长:“可妻也。” 5.名词带介宾结构做补语,这个名词活用为动词。 a.晋军(于)函陵,秦军(于)氾南。 b.唐浮图慧褒始舍于其址。 6.名词用“而”同动词或动宾词组连接时,活用为动词。 a.三代不同礼而王,五霸不同法而霸。 7.名词在“所”“者”结构中便活用为动词。 a.置人所罾鱼腹中。

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