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Abstract With ‘Victoria’ grape as a material， the effects of increasing 1 200 mg/L of CO2 on yield， soluble solid content and downy mildew of grape were studied in greenhouse. The results showed that when CO2 was increased at a rate of 1 200 mg/L in greenhouse， ‘Victoria’ fruit was harvested 8 d in advance， and compared with no increase of CO2， the transverse diameter， longitudinal diameter， average single grain weight， commodity fruit yield at early stage and total yield of commodity fruit increased by 35.07%， 7.37%， 17.39%， 24.88%， 46.14% and 20.61%， respectively， with very significant differences. The incidence and disease index of grape downy mildew decreased by more than 50%， indicating a significant control effect. Therefore， increasing CO2 in greenhouse could bring forward grape harvest， significantly improve yield and soluble solids content of grape and reduce the incidence of grape downy mildew.
Key words Increasing CO2; Production; Soluble solids; Downy mildew
CO2 is an important raw material for photosynthesis， and more than 90% of dry matter in plants is produced through photosynthesis from CO2[1]. In recent years， more and more attention has been paid to the research and extension of CO2 fertilization technology both at home and abroad. CO2 fertilization technology was studied as early as the beginning of the 19th century. Japan improved the yields of tomato， cucumber and strawberry averagely by 32.6%， 19.2%， and 20%-50%， respectively， through the application of gas fertilizer CO2[2]. Gas fertilizer CO2 was commonly applied in greenhouse in Denmark， Japan， English and America in the middle of the 20th century[3-4]， while China was beginning to start the study on facility CO2 fertilization technology until the end of the 20th century. Researches have shown that increasing greenhouse CO2 concentration is beneficial to the improvement of yield and quality of greenhouse vegetables[5-8]. However， few studies have been conducted on gas fertilizer CO2 in grape cultivation. In this experiment， with greenhouse grape variety ‘Victoria’ as an experimental material， the effects of increasing 1 200 mg/L of CO2 on yield， quality and diseases of greenhouse grape were studied， aiming at provide reference for application of gas fertilizer CO2 on greenhouse grape.
Materials and Methods
Experimental materials
The tested grape variety ‘Victoria’ was provided by Baotou Runze Garden Agricultural Science and Technology Farmers Professional Cooperative. Intelligent automatic CO2 control equipment was provided by Hesheng Group. Experimental methods
This study was carried out at Baotou Runze Garden Agricultural Science and Technology Farmers Professional Cooperative （Planting Base of the Inner Mongolia Autonomous Region Standardized Organic Grape Cultivation System Project） in 2017. The production environment of this cooperative satisfies the production standards of organic grape， and there are six greenhouses for planting ‘Victoria’ grape with a total area of 13 340 m 2， adopting Vshaped fixture cultivation mode. The plants were planted according to a spacing of 50 cm×200 cm. The experiment was designed with two treatments， i.e.， treatment A： applying CO2 1 200 mg/L， and control （CK）： no increase of CO2.
CO2 was increased at flowering stage （from March 10 to May 30）. Every greenhouse was divided to two treatments which were separated with thick nonwoven fabric plus plastic sheet. Conventional field management of water and fertilizer was adopted. Gas fertilizer CO2 was applied 0.5 h after curtain rolling. After entering May， because straw matting was not covered in the nighttime， the application of gas fertilizer CO2 was started 1 h after sunrise when the temperature inside the facility was higher than 20 ℃ and stopped 0.5 h before ventilation. In a cloudy day， the temperature in the facility was lower， CO2 was not applied to avoid CO2 poisoning.
Determination of items
Determination of grape transverse and longitudinal diameters
The transverse and longitudinal diameters of the berry were measured with a vernier caliper. Five clusters were selected from each plot， and three grains were selected from each cluster and determined， obtaining three values which were averaged.
Determination of fruit weight and soluble solid content
Average single grain weight： At maturation stage， 10 plants were randomly selected from each treatment， and two grape clusters satisfying commodity fruit requirements were selected for measurement of total weight of 100 grains， which was converted to average single grain weight.
Commodity fruit yield at early maturation stage： Commodity fruit yield within 15 d after starting harvest was calculated and converted to yield per 667 m 2.
Total yield of commodity fruit： Yield of grape actually harvested from each treatment was converted to yield per 667 m 2.
Soluble solid content： Ten fullyripe grape clusters were selected from each treatment， and 10 grains were selected from each cluster. Soluble solid contents were determined with an handheld refractometer according to Fruit Tree Germplasm Resource Descriptor[9]， and averaged. Investigation of downy mildew
The growth stages in each plot were recorded， and downy mildew condition at fruit developing stage was investigated. Downy mildew condition was graded as below.
Grade 0： Normal growth vigor， spread leaf， with no disease spots on whole leaf （fruit）; grade 1： normal growth vigor， spread leaf， with light disease spots on old leaves at the base having an area accounting for 25% or less of total leaf area; grade 2： weak growth vigor， with disease spots having an area accounting for 25%-50% （including 50%） of total leaf area， suffering from obvious disease slightly influencing yield; grade 3： very weak growth vigor， with disease spots having an area accounting for 50%-75% （including 75%） of total leaf area， in severe disease influencing yield; grade 4： very weak growth vigor， with disease spots having an area accounting for more than 75% of total leaf area， or leave withered， fruit rotted， suffering from a great yield loss.
Incidence （%）=Number of diseased plants/Total number of plants×100;
Disease index （%）=Σ（Grade of disease×Value of each grade）/（Total number of plants×Value of the highest grade）×100.
Data analysis
Data were processed with Excel， and variance analysis was performed by LSD method.
Results and Analysis
Effect of increasing CO2 on yield of greenhouse grape
It could be seen from Table 1 that treatment A （increasing CO2） showed an average transverse diameter of 28.5 mm， which was higher than the CK by 35.07%， and its longitudinal diameter was 33.5 mm， which was higher than the CK by 7.37%. Treatment A had an average single grain weight of 10.8 g， which was higher than the CK by 17.39%. It showed a fruit yield at early stage of 1 100.16 kg/667 m 2， which was higher than the CK by 24.88%. The total yield of commodity fruit in treatment A was 2 071.56 kg/667 m 2， which was higher than the CK by 46.14%. The increasing CO2 treatment showed indices all very significantly higher than the CK.
Effect of increasing CO2 on soluble solid content in greenhouse grape
It could be seen from Table 2 that treatment A had the soluble solid content of 15.8%， which was significantly improved by 20.61% compared with the CK.
Effect of increasing CO2 on downy mildew in greenhouse grape
It could be seen from Table 3 that treatment A had the morbidity of downy mildew reaching 25.07%， which was very significantly lower than the CK by 56.95%. The morbidity index was 2.01%， which was very significantly lower than the CK by 66.39%. Discussion and Conclusions
The results of this study showed that greenhouse grape applied with gas fertilizer CO2 at a rate of 1 200 mg/L reached the maturation stage on May 12， and the normal treatment without increasing CO2 reached the maturation stage on May 20. Greenhouse grape increased with gas fertilizer CO2 could enter the market 8 d in advance， with a price improved by 3 yuan/kg， so farmers income could be improved significantly.
There have been many studies on increasing gas fertilizer CO2 to facility vegetables. MADSEN[10] showed that when CO2 concentration was lower than 800 mg/L or higher than 1 500 mg/L， vegetable yield was equivalent to that under normal CO2 concentration （300 mg/L）， so a middle value， 1 200 mg/L was selected as the CO2 application concentration. Artificially increasing CO2 could remarkably enhance photosynthesis of grape leaf， improve chlorophyll content in grape leaf， thickness of grape leaf and accumulation of starch content， and accelerate the ripe speed of grape fruit， thereby finally improving single grain weight and increasing grape yield[11]. Under the condition in this experiment， increasing CO2 could bring forward crops listing date， and improve commodity fruit yield at early stage， which accords with the research report of Yao et al.[12] on tomato.
CO2 not only could provide energy desired by photosynthesis of plants， but also could serve as an important signal of improved fruit quality[13-14]. Soluble solid content in grape is an important factor of interior fruit quality， and improving soluble solid content in grape and improving grape quality are more satisfactory. Downy mildew of grape breaks out and is epidemic in Baotou area， and seriously threatens grape planting in Baotou area. Therefore， through the increase of 1 200 mg/L of gas fertilizer CO2， greenhouse grape "Victoria" showed the incidence and disease rate of downy mildew reduced by more than 50%， exhibiting very significant control effect， which meant that ordinary grape had been converted to grape reaching the standard of organic grape.
Greenhouse grape "Victoria" enter the market 8 d in advance under the increase of 1 200 mg/L of gas fertilizer CO2， and its longitudinal diameter， transverse diameter， average single grain weight， commodity fruit yield at early stage， total yield of commodity fruit and soluble solid content all increased compared with normal condition without increasing CO2， with very significant differences. And increasing CO2 effectively reduced the occurrence of downy mildew and controlled the application of agrochemicals， so the grape satisfied the standard of organic grape production. References
[1] SUN ZQ， BAI YL， ZHANG HM， et al. The effect of CO2 enrichment on photosynthetic characteristic of squash in Heliogreenhouse[J]. Journal of Henan Agricultural University， 1997， 31（4）： 375-378. （in Chinese）
[2] WANG TS. Application and countermeasure of CO2 gasfertilizer technology in sunlight greenhouses[J]. Journal of Shanxi Agricultural Sciences， 2006， 34（2）： 49-51. （in Chinese）
[3] LI YX， CHEN DK. The physiological and biological reactions of cucumber to CO2 application in greenhouse[J]. Journal of Shenyang Aricultural University， 2000， 31（1）： 86-88. （in Chinese）
[4] ZHANG Y， WANG JC， XUE QL， et al. The application of CO2 fertilizer technology in equipped environment[J]. Acta Agriculturae BorealiSinica， 2006， 21（S）： 109-113. （in Chinese）
[5] SUN Q， CUI SM， SONG Y， et al. Impact of illumination and temperature performance of blanketinside solar greenhouse and CO2 enrichment on cucumber growth and development[J]. Northern Horticulture， 2016（9）： 121-125. （in Chinese）
[6] GAO Y， CUI SM， SONG Y， et al. Effects of CO2 enrichment on growth and photosynthetic characteristics of different rootstocks grafted cucumber seedlings[J]. Inner Mongolia Agricultural Science， 2017（1）： 116-120. （in Chinese）
[7] WANG YQ， CUI SM， FANG H， et al. Influence of elevated CO2 and high temperature on the permeability of membrane and protective enzymes of grafting cucumber seedlings in the greenhouse[J]. Acta Agriculturae BorealiSinica， 2012， 27（1）： 159-160. （in Chinese）
[8] LIU JC， CUI SM， WU YF， et al. The effect of air humidity on the stress physiology of the grafted cucumber in high temperature plastic tunnel under CO2 enrichment[J]. Acta Agriculturae BorealiSinica， 2012， 29（1）： 113-118. （in Chinese）
[9] PU FS. Fruit tree germplasm resource descriptor[M]. Beijing： China Agriculture Press， 2008. （in Chinese）
[10] MADSEN E. Effect of CO2 concentration on growth and fiuit production of tomato plants[J]. Acta Agriculturae Scandinavica （Section B）， 1974（24）： 242-246.
[11] FRYDRYCH J. Factors affecting photo synthetic productivity of sweet pepper and tomato grown in CO2 enriched atmosphere[J]. Plant Physiol， 1984， 62： 95-101.
[12] YAO G， ZHANG S. Effects of CO2 application on growth and development and yield of tomato in protected area[J]. Inner Mongolia Agricultural Science and Technology， 1998（6）： 3348. （in Chinese）
[13] PIERIK R， KEUSKAMP DH， SASIDHARENS R， et al. Light quality controlsshoot elongation through regulation of multiple hormones[J]. Plant Signalling & Behavior， 2009， 4（8）： 755-756.
[14] HEIJDE M， ULM R. UVB photoreceptormediated signaling inplants[J].Trends in Plant Science， 2012（17）： 230-237.
Key words Increasing CO2; Production; Soluble solids; Downy mildew
CO2 is an important raw material for photosynthesis， and more than 90% of dry matter in plants is produced through photosynthesis from CO2[1]. In recent years， more and more attention has been paid to the research and extension of CO2 fertilization technology both at home and abroad. CO2 fertilization technology was studied as early as the beginning of the 19th century. Japan improved the yields of tomato， cucumber and strawberry averagely by 32.6%， 19.2%， and 20%-50%， respectively， through the application of gas fertilizer CO2[2]. Gas fertilizer CO2 was commonly applied in greenhouse in Denmark， Japan， English and America in the middle of the 20th century[3-4]， while China was beginning to start the study on facility CO2 fertilization technology until the end of the 20th century. Researches have shown that increasing greenhouse CO2 concentration is beneficial to the improvement of yield and quality of greenhouse vegetables[5-8]. However， few studies have been conducted on gas fertilizer CO2 in grape cultivation. In this experiment， with greenhouse grape variety ‘Victoria’ as an experimental material， the effects of increasing 1 200 mg/L of CO2 on yield， quality and diseases of greenhouse grape were studied， aiming at provide reference for application of gas fertilizer CO2 on greenhouse grape.
Materials and Methods
Experimental materials
The tested grape variety ‘Victoria’ was provided by Baotou Runze Garden Agricultural Science and Technology Farmers Professional Cooperative. Intelligent automatic CO2 control equipment was provided by Hesheng Group. Experimental methods
This study was carried out at Baotou Runze Garden Agricultural Science and Technology Farmers Professional Cooperative （Planting Base of the Inner Mongolia Autonomous Region Standardized Organic Grape Cultivation System Project） in 2017. The production environment of this cooperative satisfies the production standards of organic grape， and there are six greenhouses for planting ‘Victoria’ grape with a total area of 13 340 m 2， adopting Vshaped fixture cultivation mode. The plants were planted according to a spacing of 50 cm×200 cm. The experiment was designed with two treatments， i.e.， treatment A： applying CO2 1 200 mg/L， and control （CK）： no increase of CO2.
CO2 was increased at flowering stage （from March 10 to May 30）. Every greenhouse was divided to two treatments which were separated with thick nonwoven fabric plus plastic sheet. Conventional field management of water and fertilizer was adopted. Gas fertilizer CO2 was applied 0.5 h after curtain rolling. After entering May， because straw matting was not covered in the nighttime， the application of gas fertilizer CO2 was started 1 h after sunrise when the temperature inside the facility was higher than 20 ℃ and stopped 0.5 h before ventilation. In a cloudy day， the temperature in the facility was lower， CO2 was not applied to avoid CO2 poisoning.
Determination of items
Determination of grape transverse and longitudinal diameters
The transverse and longitudinal diameters of the berry were measured with a vernier caliper. Five clusters were selected from each plot， and three grains were selected from each cluster and determined， obtaining three values which were averaged.
Determination of fruit weight and soluble solid content
Average single grain weight： At maturation stage， 10 plants were randomly selected from each treatment， and two grape clusters satisfying commodity fruit requirements were selected for measurement of total weight of 100 grains， which was converted to average single grain weight.
Commodity fruit yield at early maturation stage： Commodity fruit yield within 15 d after starting harvest was calculated and converted to yield per 667 m 2.
Total yield of commodity fruit： Yield of grape actually harvested from each treatment was converted to yield per 667 m 2.
Soluble solid content： Ten fullyripe grape clusters were selected from each treatment， and 10 grains were selected from each cluster. Soluble solid contents were determined with an handheld refractometer according to Fruit Tree Germplasm Resource Descriptor[9]， and averaged. Investigation of downy mildew
The growth stages in each plot were recorded， and downy mildew condition at fruit developing stage was investigated. Downy mildew condition was graded as below.
Grade 0： Normal growth vigor， spread leaf， with no disease spots on whole leaf （fruit）; grade 1： normal growth vigor， spread leaf， with light disease spots on old leaves at the base having an area accounting for 25% or less of total leaf area; grade 2： weak growth vigor， with disease spots having an area accounting for 25%-50% （including 50%） of total leaf area， suffering from obvious disease slightly influencing yield; grade 3： very weak growth vigor， with disease spots having an area accounting for 50%-75% （including 75%） of total leaf area， in severe disease influencing yield; grade 4： very weak growth vigor， with disease spots having an area accounting for more than 75% of total leaf area， or leave withered， fruit rotted， suffering from a great yield loss.
Incidence （%）=Number of diseased plants/Total number of plants×100;
Disease index （%）=Σ（Grade of disease×Value of each grade）/（Total number of plants×Value of the highest grade）×100.
Data analysis
Data were processed with Excel， and variance analysis was performed by LSD method.
Results and Analysis
Effect of increasing CO2 on yield of greenhouse grape
It could be seen from Table 1 that treatment A （increasing CO2） showed an average transverse diameter of 28.5 mm， which was higher than the CK by 35.07%， and its longitudinal diameter was 33.5 mm， which was higher than the CK by 7.37%. Treatment A had an average single grain weight of 10.8 g， which was higher than the CK by 17.39%. It showed a fruit yield at early stage of 1 100.16 kg/667 m 2， which was higher than the CK by 24.88%. The total yield of commodity fruit in treatment A was 2 071.56 kg/667 m 2， which was higher than the CK by 46.14%. The increasing CO2 treatment showed indices all very significantly higher than the CK.
Effect of increasing CO2 on soluble solid content in greenhouse grape
It could be seen from Table 2 that treatment A had the soluble solid content of 15.8%， which was significantly improved by 20.61% compared with the CK.
Effect of increasing CO2 on downy mildew in greenhouse grape
It could be seen from Table 3 that treatment A had the morbidity of downy mildew reaching 25.07%， which was very significantly lower than the CK by 56.95%. The morbidity index was 2.01%， which was very significantly lower than the CK by 66.39%. Discussion and Conclusions
The results of this study showed that greenhouse grape applied with gas fertilizer CO2 at a rate of 1 200 mg/L reached the maturation stage on May 12， and the normal treatment without increasing CO2 reached the maturation stage on May 20. Greenhouse grape increased with gas fertilizer CO2 could enter the market 8 d in advance， with a price improved by 3 yuan/kg， so farmers income could be improved significantly.
There have been many studies on increasing gas fertilizer CO2 to facility vegetables. MADSEN[10] showed that when CO2 concentration was lower than 800 mg/L or higher than 1 500 mg/L， vegetable yield was equivalent to that under normal CO2 concentration （300 mg/L）， so a middle value， 1 200 mg/L was selected as the CO2 application concentration. Artificially increasing CO2 could remarkably enhance photosynthesis of grape leaf， improve chlorophyll content in grape leaf， thickness of grape leaf and accumulation of starch content， and accelerate the ripe speed of grape fruit， thereby finally improving single grain weight and increasing grape yield[11]. Under the condition in this experiment， increasing CO2 could bring forward crops listing date， and improve commodity fruit yield at early stage， which accords with the research report of Yao et al.[12] on tomato.
CO2 not only could provide energy desired by photosynthesis of plants， but also could serve as an important signal of improved fruit quality[13-14]. Soluble solid content in grape is an important factor of interior fruit quality， and improving soluble solid content in grape and improving grape quality are more satisfactory. Downy mildew of grape breaks out and is epidemic in Baotou area， and seriously threatens grape planting in Baotou area. Therefore， through the increase of 1 200 mg/L of gas fertilizer CO2， greenhouse grape "Victoria" showed the incidence and disease rate of downy mildew reduced by more than 50%， exhibiting very significant control effect， which meant that ordinary grape had been converted to grape reaching the standard of organic grape.
Greenhouse grape "Victoria" enter the market 8 d in advance under the increase of 1 200 mg/L of gas fertilizer CO2， and its longitudinal diameter， transverse diameter， average single grain weight， commodity fruit yield at early stage， total yield of commodity fruit and soluble solid content all increased compared with normal condition without increasing CO2， with very significant differences. And increasing CO2 effectively reduced the occurrence of downy mildew and controlled the application of agrochemicals， so the grape satisfied the standard of organic grape production. References
[1] SUN ZQ， BAI YL， ZHANG HM， et al. The effect of CO2 enrichment on photosynthetic characteristic of squash in Heliogreenhouse[J]. Journal of Henan Agricultural University， 1997， 31（4）： 375-378. （in Chinese）
[2] WANG TS. Application and countermeasure of CO2 gasfertilizer technology in sunlight greenhouses[J]. Journal of Shanxi Agricultural Sciences， 2006， 34（2）： 49-51. （in Chinese）
[3] LI YX， CHEN DK. The physiological and biological reactions of cucumber to CO2 application in greenhouse[J]. Journal of Shenyang Aricultural University， 2000， 31（1）： 86-88. （in Chinese）
[4] ZHANG Y， WANG JC， XUE QL， et al. The application of CO2 fertilizer technology in equipped environment[J]. Acta Agriculturae BorealiSinica， 2006， 21（S）： 109-113. （in Chinese）
[5] SUN Q， CUI SM， SONG Y， et al. Impact of illumination and temperature performance of blanketinside solar greenhouse and CO2 enrichment on cucumber growth and development[J]. Northern Horticulture， 2016（9）： 121-125. （in Chinese）
[6] GAO Y， CUI SM， SONG Y， et al. Effects of CO2 enrichment on growth and photosynthetic characteristics of different rootstocks grafted cucumber seedlings[J]. Inner Mongolia Agricultural Science， 2017（1）： 116-120. （in Chinese）
[7] WANG YQ， CUI SM， FANG H， et al. Influence of elevated CO2 and high temperature on the permeability of membrane and protective enzymes of grafting cucumber seedlings in the greenhouse[J]. Acta Agriculturae BorealiSinica， 2012， 27（1）： 159-160. （in Chinese）
[8] LIU JC， CUI SM， WU YF， et al. The effect of air humidity on the stress physiology of the grafted cucumber in high temperature plastic tunnel under CO2 enrichment[J]. Acta Agriculturae BorealiSinica， 2012， 29（1）： 113-118. （in Chinese）
[9] PU FS. Fruit tree germplasm resource descriptor[M]. Beijing： China Agriculture Press， 2008. （in Chinese）
[10] MADSEN E. Effect of CO2 concentration on growth and fiuit production of tomato plants[J]. Acta Agriculturae Scandinavica （Section B）， 1974（24）： 242-246.
[11] FRYDRYCH J. Factors affecting photo synthetic productivity of sweet pepper and tomato grown in CO2 enriched atmosphere[J]. Plant Physiol， 1984， 62： 95-101.
[12] YAO G， ZHANG S. Effects of CO2 application on growth and development and yield of tomato in protected area[J]. Inner Mongolia Agricultural Science and Technology， 1998（6）： 3348. （in Chinese）
[13] PIERIK R， KEUSKAMP DH， SASIDHARENS R， et al. Light quality controlsshoot elongation through regulation of multiple hormones[J]. Plant Signalling & Behavior， 2009， 4（8）： 755-756.
[14] HEIJDE M， ULM R. UVB photoreceptormediated signaling inplants[J].Trends in Plant Science， 2012（17）： 230-237.