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2024-09-26 PL-INT-2024-008813 GP 2045 Draft Comments - A. Longo
DeVera, Ashley From: Angela Longo < Sent: Thursday, September 26, 2024 8:37 AM To: Planning General Plan Subject: Critique of Hawaii general plan Aloha I am Dr Angela Longo with PhD in Biochemistry and an OMD an energy medicine and I know for a fact that we need every molecule of CO2 in the atmosphere to produce the food needed by the world. It is unscientific to believe that mankind produces too much. I have thousands of signatures of scientists who know that this argument does not stand. The weather change is being caused by a natural phenomena called the precession of the Earth's axis. Any scientists who studies the earth learns this in high school or should if they're paying attention. We can only protect the people on the coast since this is inevitable natural change. These plans to sequester people does not make sense and is unnecessary and unhealthy for the physical and mental well-being of our people. Please rewrite your plan to give people more space to live and grow the necessary food that we need to live on. I am quoting one of my friends who research the numbers. From Donna thompson: Hawaii General plan mentions the potential to sequester CO2 as rationale to reduce man made impacts, even though temperature levels are drastically lower now than they have been in 485 million years (see chart above) Photosynthesis activity in plants requires CO2 and only makes up 0.04% of our atmosphere. Reduction of this necessary element would have a catastrophic chain reaction and is a risk to ALL carbon based lifeforms. Several long-term studies have provided strong support demonstrating that growing crops under elevated [CO2] can increase photosynthesis and result in an increase in yield, flavour and nutritional content (including but not limited to Vitamins C, E and pro-vitamin A). In the case of tomato, increases in yield by as much as 80% are observed when plants are cultivated at woo ppm [CO2], which is consistent with current commercial greenhouse production methods in the tomato fruit industry. These results provide a clear demonstration of the potential for elevating [CO2] for improving yield and quality in greenhouse crops. https://academic.oup.com/hr/article/10/4/uhad026/7049409?login=false This testimony is to provide overall big picture of extinction level event(s) to instigate community emergency preparedness as part of the overall plan Increased solar radiation as a result of the earths weakening magnetosphere and polarity shift will significantly impact survivability of our community. There will be significant disruptions of the supply chain, challenges with food production, energy blackouts, and limited communications. Community preparedness for such events are not documented in the draft general plan, meanwhile many eastern hemisphere nations including Russia, Thailand, and Japan have been preparing for these events with local civil defense for years. Citizens of Hawaii county are aware of the influx of billionaires over the last 20 years that have had impact on county policy and planning. Many have taken permanent residence and hedging Hawaii island has a good chance of survivability during these changes. In light of this information, my suggestion is for our county to birth its own plan based on the needs of the community expressed in numerus testimonies and include space weather scenarios as potential hazards. County officials would be wise to form a committee of experienced personnel from various departments - including the local community members with relevant capabilities that can prioritize and draft a community plan that includes emergency preparedness that is relevant to our our island. We need to focus on independence with agricultural food production, innovative energy and building technologies and resilience thru preparation to create our own future we want to see. Dr Angela Longo living in Kamuela Hawaii 2 JOURNAL ARTICLE Feeding the world: impacts of elevated [CO2] on nutrient content of greenhouse grown fruit crops and options for future yield gains ,,,, Horticulture Research,Volume 10,Issue 4,April 2023,uhadO26,https://doi.org/10.1093/hr/uhadO26 Published: 21 February 2023 Article history v Abstract Several long-term studies have provided strong support demonstrating that growing crops under elevated[CO2]can increase photosynthesis and result in an increase in yield,flavour and nutritional content(including but not limited to Vitamins C,E and pro-vitamin A).In the case of tomato,increases in yield by as much as 80%are observed when plants are cultivated at moo ppm[CO2],which is consistent with current commercial greenhouse production methods in the tomato fruit industry.These results provide a clear demonstration of the potential for elevating[CO2]for improving yield and quality in greenhouse crops.The major focus of this review is to bring together 5o years of observations evaluating the impact of elevated[CO2]on fruit yield and fruit nutritional quality.In the final section,we consider the need to engineer improvements to photosynthesis and nitrogen assimilation to allow plants to take greater advantage of elevated CO2 growth conditions. Issue Section: Review Article Introduction Elevated[CO2](e[CO2])has been shown to significantly improved light saturated photosynthetic carbon assimilation rates(ASat)by increasing the efficiency of Rubisco CO2 assimilation(carboxylation)over the alternate RuBP oxygenation(02 assimilation),which results in enhanced growth and yield[i,2](Figure 1). Small Large elevation in elevation in Ico,l [t0,1 Ambient[CO,i 420 ppm Rubisco rate As 100A surpasses 550 Since Rubisco already limiting In CRC. ppm A,.is limited by the saturated,further leading to reduced rate of RuBP regeneration. increase in yield not carbon assimilation Rubisco no longer rate necessarily seen and plant biomass limiting 1 11fIncreased carbon Increased assimailtion and Increased fertilisation biomass fertilisation IIs there art optimal [CO,l to increase yield without compromising quality? Figure 1 Schematic representation of elevated[CO2]on carbon assimilation.Created with BioRender.com PDF Helps',�W Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. Fruit Effects of elevated[COO on fruit yield and quality -n.to ussm<..a..:n van yr".ra tremeata le[<0,1•43low.-120001.) 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O.Or,wromns •I.6:4..r,.,,..:n..,.. rollY[O}r,rarmenrr ra60aar3-WO worn) •}to a5' -. .y4<au.k.00u.Mxrore)breiCO,1:,....oom. -Ur . .133%increase wet.fUelin.l:11werx f0}% IIE e,16sD reponeem(r00'Y•9wpam� iou,'errna C. m011.. roD <•.01.edreas.1 e.Dee �[a-/aroma<arn�lwnn(eters,uar:M.l.naooll.n'reaus tlepv.uwn:n<re F OIIw Irmrr[.[Med la9q,69q,9Sg9om7. :w spec**pre of CO,f.tLr^'9oaara[on be 00[0..9000 as.0.10,.0c.,.n<r:,r:.,•rr:0Of()O.>ASO an f w! dua';ryWve.09.0.tot.GreM:et CO,frnmsoaan op9earslo 9•.09[,nr.;,r.r:oor..1o,„„ry r ..„:mr<m ,..>nr::r«0.'r .,. ,., 0).,0.0.0 :ronan ate ro-s%w.tar0a•.rrw o ...Oran,.0:a.,.,o moo.rn, I coal 1 Iva Lr,9: xd Now waarourxn rlx wnw Oder'ro tion,05)00 and M5. aa..red,orood.aa'a.wrnx yew and O0aney Figure 2 Effects of elevated[CO2]on yield and quality of fruiting crops.Created with BioRendercom The majority of research evaluating the impact of e[CO2]on fruit crop production has been carried out in controlled environment conditions (chambers),polytunnels and commercial greenhouses where crops are grown in e[CO2],and focus almost exclusively on soft fruit such as strawberry,tomato and cucumber.Early work in the 1980's suggested that e[CO2]increased the average yield of all plants tested by approximately 30/0,with optional[CO2]concentration for growth and yield in the range of 700 to 90o ppm with concentration in excess of loon ppm having a negative impact on plant growth and yield[3-6].In the case of vegetable cops,much of the work has been carried out in controlled environments, in which elevated[CO2](800-90o ppm)increased lettuce,carrot,and parsley yield by 18%,19%,and 17%,respectively in greenhouse grown crops. However,the yields of leek,chinese cabbage and celery were not significantly affected by increases in growth[CO2]concentration[7].A meta- analysis of 107 selected articles showed that e[CO2]results in an increase in vegetable number(yield)by on average 32%and vegetable mass by 11°/u [8].Furthermore,a meta-analysis of 57 articles consisting of 1015 observations found that e[CO2]has both positive and negative impacts on vegetable quality.For example,whilst concentrations of fructose(+14.2°/0),glucose(+13.2%),total soluble sugar(+17.5%),total antioxidant capacity(+59.0%),total phenols(+8.9%),total flavonoids(45.5%),vitamin C(+9.5%),and calcium(+8.2%)increased in the edible part of vegetables,protein(-9.5%,)nitrate(-18.o%),magnesium(-9.2%),iron(-16.0%),and zinc(-9.4%)decreased[9].Moreover,a meta-analysis of legumes found a reduction in zinc and iron(and in non-legumes a reduction in protein)when plants were grown under e[CO2](see Myers et al[10]). In 2018,Zhu et al[11]confirmed these results,and moreover demonstrated that rice grown under e[CO2]showed consistent declines in the quantities of vitamins Bi,B2,B5,and B9 and,an increase in vitamin E.Finally,studies have shown that grains(wheat,rice,and barley),legumes, and maize-have a 4-1o%reduction in iron concentrations of when grown under e[CO2](-55o ppm)[12].These results shown that e[CO2]can positively and negatively impact on legumes,grain and vegetables on a crop-by-crop basis and simultaneously alter quality attributes in the same harvestable material. The aim of this review is to provide an overview of the current available data of the impact of elevated[CO2]on fruiting crops production in commercial growing systems.This paper examines these studies and the long-term implications of e[CO2]on the yield and quality of fruit required to feed a growing population.In the last section,we discuss the potential for designing crops for these new growing environments and allowing them to take full advantage of the introduced CO2,potentially increasing crop yield,reducing costs for commercial producers,and improving quality of the final product providing high nutritional value to consumers. Impact of elevated [CO2] on yield and quality of GREENHOSUE grown crops Impact of elevated [CO2]on solanaceous crops Commercially,tomato crops are grown in greenhouses with e[CO2],in some cases as high as 2000 ppm.The effects of e[CO2]of fruit yield and quality has been extensively studied(Figure 2).Under e[CO2],tomato fruit yield increases ranged from 7%—125%with[CO2]ranged from 45o ppm —1200 ppm compared with plants grown under a[CO2].An increase in the quantity of non-reducing sugars(glucose and fructose)has been reported [13-17]and fully ripe tomatoes grown in an e[CO2]were found to be preferable for consumption in sensory panels[13].As liking sweetness has been shown to be a universal trait[18],it is possible that this increase in sugar is responsible for preference of the carbon enriched tomato fruits.An increase in vitamin C was also found between most studies[13,15,16,19],potentially improving the health benefit gains from consumption of carbon-enriched grown tomatoes(Table 1).Vitamin C is an important dietary requirement and at high concentrations it has been used as a treatment for cancer,arteriosclerosis,and cardiovascular diseases[2o-22].These results suggest that increasing environmental[CO2]could contribute to an increase in Vitamin C improving their nutritional value for the consumer.However,growth at e[CO2]does not have the same impact on all species,as another studies in barley reported a significant decrease in Vitamin C content[23]highlighting the species—species response differences to e[CO2]and suggesting that high carbon growth environments may not always provide the best outcome for the consumer PDF even though increases in yield maybe the producers primary concern(see Fenech et al.[24]and references therein). Hep([ Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. level of CO2 enrichment impacts fruit quality and careful consideration is needed to ensure an appropriate balance between levels of e[CO2]and final yield.Lycopene is an important phytonutrient,is sold commercially as a dietary supplement,and has been reported to possess anti-cancer properties and can improve cardiovascular health[26,27]. I-carotene is the precursor for Vitamin A,also known as retinol.Vitamin A is an essential micronutrient playing important roles in growth and development,vision[281 and the immune system[29].More than a third of all pre-school children and a significant number of pregnant women around the world are affected by Vitamin A deficiency,increasing the risk of night blindness and miscarriage[3o,31].Importantly,most people suffering from a deficiency in Vitamin A show no clinical symptoms resulting in a phenomenon termed"Hidden Hunger"[32].Production of crops with increased Vitamin A is therefore an important target for improving the diet and health of these at-risk groups;enhanced uptake of carbon may be a useful approach to achieve this.Increases in the Vitamin A precursor(3-carotene has been observed in tomato fruit grown under e[CO2]of 800- 900 ppm,in addition to a 28°7°increase in vitamin C at ripe stage and an-8%increase in total soluble solids(Table 1).Zhang et al.[13],suggested that under these growth conditions,improved vitamin A and C and increased carotenoid content may be attainable. Carotenoids are also the precursors of several flavour and aroma compounds.I-carotene is cleaved by carotenoid cleavage dioxygenases CCDM and CCD4[26,33-35],to form the aromatic apocarotenoid 0-ionone,which is important to tomato fruit flavour.Furthermore,lycopene,shown to increase under e[CO2]is cleaved by CCD1 to form several important flavour and aroma compounds including 6,10-dimethyl-3,5,9-undecatrien-2- one(pseudoionone[34];,6-methyl-5-hepten-2-one(MHO[36];and geranial[37].MHO has been shown to be an important contributor to tomato fruit flavour[38,39]and has also been shown to accumulate in tomato fruit with higher lycopene levels[40].It is therefore apparent that growth in e[CO2]can increase a range of key flavour and nutraceutical precursor compounds present in tomato fruit;this phenomenon deserves further study, the optimal levels of[CO2]are currently not clear and more work is needed to better understand the relationship between CO2,assimilation carotenoid content,flavour and overall quality(Table 2). Similar results have also been found in pepper crops,with yield increase of 12.9%—370.2%reported when grown at e[C07]of between 450 ppm— l000 ppm(Table 2)with most other studies reporting yield increases in the range of 12.9%—47.4%in the absence of other parameters[41-47]. However,it should be noted that growth at-800 ppm e[CO2]was found to reduce sweet pepper total amino acid content by up to 29%,including reductions in the sweet tasting amino acids alanine and glycine,which could be detrimental to the perceived fruit flavour[42].Yield was also found to vary with different irrigation programmes[41,48],nitrogen sources[48],substrate salinity[42,44]and pruning regimens[46].Given that previous work in tomato has shown an increase in potential phytonutrients in fruit grown at 55o ppm and a decrease in those grown at 700 ppm, further research is needed to better identify the specific quantity of CO2 fertilisation necessary for maximally improved yield in solanaceous crops, especially when considering that CO2 uplift is often accompanied by additional treatments,such as increased nutrient and nitrogen fertilisation (Figure 2). Impact of elevated atmospheric[CO2]on yield and nutritional quality of tomato Table 1 Impact of elevated atmospheric[CO2]on yield and nutritional quality of tomato CO2Treatment AdditionalTreatment(s) Fruit Yield Fruit Quality Ref 510 ppm N/A 9.9%increase in fruit yield. N/A [45] 590 ppm root drying Fruit dry weight not significantly affected by[CO2] N/A [15 across all irrigation treatments. 5] 375 ppm- Ozone treatment 24%increase in fruit yield.31%decrease in fruit yield N/A [15 675 ppm 80 nmol mol when exposed to ozone.Ozone and CO2 treated fruit 6] yields were not significantly different to plants grown in ambient conditions. 550 ppm N/A 54%increase in fruit yield at 550 ppm and 125% 1.4%-11.4%decrease in total soluble solids,27.3%- [19] 700 ppm increase in fruit yield at 700 ppm. 31.8%decrease in total acids and 16.1%-29.0% increase in vitamin C. +2°C increase in 18.4%-21.4%increase in fruit yield due to increased 10%increase in total sugars,44%increase in vitamin C, [25] temperature [CO2]. 32%increase in lycopene at e[CO2]in absence of other treatments.e[CO2]rescues reduction in quality from increased temperature. 650 ppm N/A 17%increase in fruit yield at 650 ppm and 48% N/A [3] 1000 ppm increase in fruit yield at 1000 ppm. 700 ppm Doubled N fertilisation N/A 13%-25%decrease in fruit lycopene content across [15 harvests with e[CO2].9%increase in fruit lycopene 7] content with increased N fertilisation. UV-B exposure up to 38%increase in fruit yield in absence of additional UV- Up to-22%increase in soluble sugars,-24%increase in [15] 1.744 kJ m-2 B treatment,up to 46%increase in fruit yield with UV-B organic acids,-40 increase in vitamin C and-47% treatment. increase in lycopene content of fruits grown under e[CO2]and UV-B treatment. PDF 700 ppm N/A -30%increase in individual fruit weight. -18%increase in vitamin C.-Up to 20%reduction in [:6] H el rsly, 900 ppm major acids(citric,malic,oxalic).-45%increase in Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. 800 ppm 0-0.5 g N kg-1 soil.Soil Across all treatments,-3.3%-28%increase in total -17.9%-11.9%increase in total fruit sugars and-18.9% [15 water content 25%-35% fruit yield. -12.7%increase in total fruit acids across all treatments. 9] Saliniti treatments at 5-7 13%increase in yield in carbon-enriched atmosphere 7%increase in total soluble solids.No significant change [14] dS m and 31%reduction in marketable fruit yield in in citric acid content.Organoleptic qualities of tomatoes increased salinity. grown under increased salinity and CO2 found preferable in sensory trials. 100 or 200 mg N kg 1 soil, 8°/o increase in fresh fruit yield with increased[CO2]. No significant difference in total sugars,organic acid or [16 70%irrigation of control fruit firmness for fruits grown in e[CO2]. 0] and root drying (Continued) Continued Table 1 Continued CO2Treatment Additional Fruit Yield Fruit Quality Ref Treatment(s) 800 ppm- N/A N/A -28%increase in vitamin C at ripe stage,-8%increase in total soluble [13] 900 ppm solids and no difference in total acids.Marked preference in sensory trials for fruits grown under enriched[CO2]. 900 ppm N/A 30%increase in marketable fruit yield. N/A [16 1] 100 limo!s-1 m-2 12%-15%increase in yield under supp N/A [16 supp lighting lighting,7%increase in yield in absence of 2] additional treatment. N/A 22%increase in total fruit yield for plants N/A [16 grown in e[CO2]. 3] 1000 ppm N/A 30%increase in total fresh fruit yield per N/A [16 plant. 4] 43%increase in total fruit yield. No significant effect on fruit quality parameters. [16 5] 74.3%-83.6%increase in tomato fresh 16.1%-20.9%increase in total sugars.20.0%-24.7%decrease in [17] weight per plant. vitamin C.4.79%-6.8%decrease in total acids. 15.6%increase in fruit yield across 8 N/A [16 different cultivars. 6] 1200 ppm Salinity up to4.58x >40%loss in dry fruit yield at highest Increased salinity and[CO2]combined increases total sugar and acid [16 control salinity treatment completely offset by content by up to-30%. 7] increased[CO2]. In chili pepper,yield increases of 43.80/0—142%were reported for e[CO2](in the range of 50o ppm—1140 ppm).These yield increases were in part attributed to an increase in the size of fruits[49].However,in controlled environments a 4°C increase in temperature decreased yield,even at e[CO2](75o ppm)[5o,51],indicating that carbon enrichment is not sufficient to rescue yield where glasshouse facilities or growth tunnels experience periods of elevated temperature in an extreme climate change scenario.Carbon-enriched growth was found to increase the capsaicinoid content of fruits,resulting in an increase in Scoville Heat Units(SHU)[49,52].This approach therefore has potential for producing hotter varieties of chili,a growing and competitive market.However,at the same time Vitamin C concentration decrease by up to 15.84%[53],reducing potential health benefits gained from growing chilli plants under e[CO2].(Table 2). These reports suggest that the effects of growing crops in e[CO2]can have both a positive influence on yield and nutritional quality,however, growth at[CO2]levels above what is optimum can negatively impact some quality traits. Impact of elevated [CO2] on rosaceous crops Rosaceous crop research in this area has focused primarily on cultivated strawberry with a small number of studies on raspberry and Nashi pear (Table 3).This is likely due to the relatively smaller size and rapid growth of strawberry compared to other commercially important rosaceous fruit species,such as tree fruits,like apple and cherry,and woody stemmed shrub fruits,like raspberry and blackberry.This makes strawberry a convenient plant to study as a rosaceous model.Furthermore,greater production of strawberry fruits would not only increase profits for growers PDF but also decrease costs for consumers,increasing the availability of healthier options.Better access to such products through economic growth is strongly correlated to reduced micronutrient malnutrition or"hidden hunger"[54]. Heirs Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. were upregulated in strawberry plants grown in an enriched-carbon atmosphere,with 14 of these being photosynthetic genes[6o],suggesting that plants respond to these atmospheric increases by increasing their ability to assimilate the excess carbon. Additional annual yield increases could be achieved by a two-week reduction in time to fruiting for plants grown in an enriched-carbon atmosphere [58,61]increasing the field season and the period of productive(fruit)growth.Several fruit quality traits are also improved by growth at e[CO2]; increases in reducing sugars,and therefore sweetness index,were reported[62,63]alongside reductions in organic acids[62].These increases in sugar-acid ratio is highly favourable for a more pleasant perception of strawberry flavour by the consumer[18]and an increase in key volatile organic compounds,including furaneol,linalool and major esters,was also reported,further enhancing the"strawberry"aroma[62].Growth in a carbon-enriched atmosphere therefore strongly enhances strawberry flavour and increases vitamin C(an important nutritional compound)by up to 13.3%alongside other antioxidant compounds[64,65].Growth in carbon-enriched atmospheres therefore simultaneously improves yield, flavour and health benefits of strawberry fruits,creating enormous potential for strategies involving enhanced photosynthesis of strawberry plants,including genetic manipulation.The greatest reported increase in fresh fruit yield where obtained when[CO2]was kept between 60o ppm— moo ppm[58],linking greater carbon assimilation to increased fresh fruit yield in strawberry and demonstrating an optimal degree of CO2 fertilisation for strawberries(Table 3). Impact of elevated atmospheric[CO2]on yield and nutritional quality of other Solanaceous crops Table 2 Impact of elevated atmospheric[CO2]on yield and nutritional quality of other Solanaceous crops Crop CO2Treatment Additional Treatment(s) Fruit Yield Fruit Quality Ref Sweet 350 ppm N/A 12.9%increase in fruit yield 350 ppm and 47.4%increase in fruit N/A [45] Pepper 450 ppm yield 450 ppm. 400 ppm 20 mmol L-1 NaCI,foliar 18.9%to 26.6%increase in yield at 400 and 800 ppm Little significant effect of increased [42] 800 ppm calcium treatment respectively.Foliar calcium treatment had no impact on yield. [CO2]on fruit inorganic nutrients or e[CO2]rescued total yield loss from high salinity. colour. 700 ppm High/low irrigation and N Fruit yield fore[CO2]increased with irrigation with no significant N/A [48] treatments difference in fruit yield at lowest irrigation. 700 ppm- N/A 18%-22%increase in total fruit yield. N/A [46] 750 ppm 800 ppm Nitrogen source and 8%and 22%increase in marketable fruit yield under salinity N/A [43, saline treatment(8 and stress and unstressed respectively.23%and 29%maximum 44,1 25 mM NaCI) increase in daily fruit harvest yield fort different nitrogen 68] sources at low salinity. 900 ppm N/A 7%increase in early yielding fruits,no change in total fruit yield. N/A [162 367 ppm- Range of irrigation Yield increased with irrigation and carbon dioxide with a N/A [41] 1000 ppm regimens maximum yield increase with both treatments of 264%. 1000 ppm N/A 51%-370%increase in fruit weight per plant. N/A [169 10000 ppm N/A 20%increase in fruit yield. N/A [61] Chili 380 ppm- +2°C and+4°C Up to 41.9%increase in fruit diameter under both increased 27%-44%increase in capsaicin [52] pepper 750 ppm temperature elevation carbon dioxide and increased temperature. across all treatments for 2 cultivars across 2 growth years. 380 ppm- +2°C and+4°C 53.8%increase in fruit number at[CO2]=550 ppm and ambient N/A [50, 750 ppm temperature elevation +2°C temperature,12.3%decrease in fruit number per plant for 51] [CO2]=750 ppm and ambient+4°C temperature.Up to-140% increase in fruit yield per plant for[CO2]=550 ppm and ambient +2°C temperature,up to-36%reduction in fruit yield per plant for[CO2]=750 ppm and ambient+4°C temperature. 380 ppm- N/A Up to 88.5%increase in number of fruits per plant,up to 13.0% No change in colour of ripe fruits.Up [49, 1140 ppm increase in fruit length,up to 15.0%increase in fruit width and to 28.6%increase in capsaicinoids in 170] up to 14.3%increase in pericarp thickness.Up to 142%increase ripe fruit. in fruit yield. 400 ppm- Natural light 92%-113%increase in total fruit yield per plant across all CO2 2%-10%decrease in soluble [171 900 ppm (233 pmol m-2 S-1)and treatments with supplementary lighting relative to ambient sugars.13%-34%decrease in supplementary light control at 400 ppm.47%-113%increase in total fruit yield per vitamin C in e[CO2].61%increase in (463 pmol m s ) plant across all CO2 treatments with natural lighting relative to capsaicin at[CO2]=550 ppm,49%- ambient control at 400 ppm. 61%decrease in capsaicin for[CO2] PDF >550 ppm. Help. Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. Tablet Continued Crop CO2Treatment Additional Fruit Yield Fruit Quality Ref Treatment(s) 1000 ppm N/A 43.80%-59.55%increase in Up to-150/0 increase in total fruit sugars.11.84%-15.84%decrease in fruit [53] fruit fresh weight per plant vitamin C,non-significant decrease in fruit titratable acids.Variable effects on across 5 cultivars. inorganic nutrient concentrations.Fruit amino acids and fatty acids mostly reduced. Eggplant 200 ppm- N/A 209%increase in fruit fresh N/A [17 3000 ppm weight and 134%increase 2] in fruit dry weight. 1000 ppm N/A 31%increase in fruit yield N/A [16 across a full year of 9] harvests. 663 ppm N/A 23.6%increase in fruit yield. N/A [45] Impact of elevated atmospheric[CO2]on yield and nutritional quality of Rosaceous crops Table 3 Impact of elevated atmospheric[CO2]on yield and nutritional quality of Rosaceous crops Crop CO2Treatment Additional Fruit Yield Fruit Quality Ref Treatment(s) Strawberry 553 ppm Nitrate treatment(4x 42%increase in fresh fruit yield in e[CO2]at high N/A [17 10 2-0 mM) N,17°/o increase in fresh fruit yield in e[CO2]at 3] low N. 400 ppm, Ambient temperature 9.9%-33.4%increase in total fruit yield at Total fruit polyphenolic content,flavonoid content, [56 650 ppm and (25°C)and elevated ambient temperature for cultivar"Albion",0.9% monomeric anthocyanin content and antioxidant ] 900 ppm (30°C) -31.2%decrease in total fruit yield at ambient content increased in correlation with e[CO2]at [65 temperature for cultivar"San Andreas". both temperatures for multiple cultivars(-9%- Elevated[CO2]rescues yield loss from elevated -325%increase overall increase at[CO2)= temperature. 900 ppm). 720 ppm 5°C increase in -120%increase in total fruit dry weight in 48%,21%,36%and 18%decrease in fruit [63 temperature,nitrate e[CO2[,-73%increase in total fruit dry weight in anthocyanin content,total phenolic content,total ] treatment(50 mL 0.1% e[CO2]with nitrogen treatment.No significant flavonoid content and total antioxidant content NH4NO3 twice per week) change in fruit yield for all other treatments respectively at e[CO2].29°/o and 35%increase in individually and in combination. fruit fructose and glucose respectively.43% increase in total sugars. 600 ppm- N/A 62%increase in total fruit yield in e[CO2]. N/A [58 1000 ppm 700 ppm- N/A 17.6%and 38.5%increase in individual fruit 7.0%-25.9%increase in glucose,fructose and [62 1000 ppm weight at +300 ppm and[CO2]= sucrose.5.2%-47.4%decrease in citric,malic and ] ambient+600 ppm respectively. quinic acids.Stepwise increase in concentration of most key volatile esters and up to 115.0%and 149.6%increase in fruit furaneol and linalool content. 700 ppm- N/A 13.3%increase in fruit ascorbic acid.Stepwise [64 1000 ppm increase in antioxidant and flavonoid compounds ] with increasing carbon dioxide. 700 ppm- 5.4%and 12.7%increase in marketable fruit N/A [15 1000 ppm yield for cultivars"Irvine"and"Chandler" 8] respectively. 1000 ppm N/A 47%increase in fruit number per plant,no N/A [55 significant change in individual fruit weight. 900 ppm, N/A 31%,43%and 51%increase in fruit yield at N/A [61 1500 ppm, 900 ppm,1500 ppm and 3000 ppm respectively. 3000 ppm Raspberry 436 ppm N/A 12%increase in total berry yield and 5% N/A [17 increase in average individual berry weight. 4] Nashi Pear 700 ppm Ambient+4°C 16.6%increase in fruit weight with e[CO2]. Up to 15.9%reduction in fruit firmness with e[CO2]. [75 PDF temperature Elevated[CO2]rescues yield loss from increased Up to 22.5%increase in total soluble solids with no ] significant change in aciditywith e[CO] Heiv temperature. g z Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. Impact of elevated [CO2]on cucurbitaceous crops Cucumber is the most studied fruit crop of the cucurbitaceae in relation to growth in carbon-enriched atmospheres(Figure z;Table 4).Improved carbon assimilation rates of up to 99%and 112%have been reported for cucumber and melon respectively when grown in e[CO2][66,67], demonstrating that growth in e[CO2]improves photosynthesis of cucurbitaceous crops. Impact of elevated atmospheric[CO2]on yield and nutritional quality of Cucurbitaceous crops Table 4 Impact of elevated atmospheric[CO2]on yield and nutritional quality of Cucurbitaceous crops Crop CO2Treatment Additional Fruit Yield Fruit Quality Ref Treatment(s) Cucumber 400 ppm, 2 mmol L-1 Up to 73%increase in fresh fruit yield for 75%increase in fruit fructose,73%increase in glucose at [66 625 ppm, 7 mmol L-1 plants grown at highest[CO2]versus plants 7 mmol L-1 at highest[CO2].No significant change in fruit ] - 1200 ppm 14 mmol r 1 NO3 grown at lowest[CO2]at greatest N titratable acidity.e[CO2]reduced dietary fibre by 13%-18% [68 fertilisation.No significant difference in across all fertilisation treatments.Up to 84%reduction in ] yield for lower N fertilisation. fruit nitrogenous compounds in e[CO2]across all nitrogen treatments. 400 ppm, 0.06 g N kg 1 soil 31%-37%increase in fresh fruit yield for Across both nitrogen treatments at[CO2]=1200 ppm,fruit [70 800 ppm, (low N), [CO2[=800 ppm and 1200 ppm at low N. fructose was increased by 5%-6%,fruit glucose was ] 1200 ppm 0.24 g N kg-1 soil 71%-106%increase in fresh fruit yield for increased by 10%-12%and starch was increased by 29%- (high N) [CO2]=800 ppm and 1200 ppm at high N 40%. 364 ppm, N/A Up to 10.2%increase in individual fruit No significant change in fruit dry matter content [45 620 ppm weight for August production in e[CO2] ] 400-500 ppm N/A 19%increase in fresh fruit yield at e[CO2] N/A [17 5] 600-700 ppm N/A 20%increase in fresh fruit yield at e[CO2] N/A [17 6] 700 ppm N/A 14.2%-18.4%increase in fresh fruit yield at Overall reduction in fruit inorganic nutrients(N,P,K,Ca,Mg). [71 e[CO2]across two crop cycles. ] 780 ppm N/A 35%increase in fresh fruit yield in N/A [17 greenhouse supplemented with[CO2] 7] versus control greenhouse. 700 ppm- N/A 20%-30%increase in marketable fruit N/A [15 1000 ppm yield across two growing seasons. 8] 900-1000 ppm 0.6°C-1.8% 35.4%increase in dry fruit mass in cooled N/A [17 cooling and e[CO2]conditions 8] 1000 ppm N/A 8.9%increase in fruit weight but no N/A [17 significant change in fruit number at e[CO2] 9] 900 ppm, N/A 18.4%-26.3%increase in fresh fruit yield N/A [61 1500 ppm, across all CO2 elevations. ] 3000 ppm Melon 400 ppm, 0,25,50 mmol NaCI Up to 29%increase in fruit yield in all N/A [67 800 ppm, e[CO2]at no additional salinity.Elevated ] 1200 ppm [CO2]partially rescues yield loss from salinity(by up to 18%)but is insufficient to fully mitigate yield loss. 1000 ppm N/A 13%increase in muskmelon fruit number N/A [16 and 8%increase in muskmelon fruit weight 9] during summer production under e[CO2] Squash 700-1000 ppm N/A 15.5%-19.7%increase in total marketable N/A [15 yield across 2 growing seasons. 8] In cucumber(Cucumis sativus),fruit yield increases for plants grown in enriched-carbon atmospheres([CO2]=45o ppm-300o ppm)ranged between 16.2%and 41%in the absence of other parameters that could alter fruit yield.In high nitrogen supplemented fertilisation,fruit yield was as high as 106%when grown under e[CO2]of 80o ppm[68],indicating the potential of increased nitrogen fertilisation alongside[CO2]enrichment to unlock the greatest yield increases in cucumber.Interestingly,when grown under e[CO2]of 1200 ppm with the addition of high nitrogen PDF fertilisation treatment,studies found a yield increase between 71%-73%[66,68],which was lower than the 106%for plants grown at e[CO2]of 80o ppm.Concentrations of[CO2]above optimal reduced stomatal density,stomatal conductance(gs),the maximum carboxylation rate(Vcmax) HeiA"' .. .. ... . . - - .. .. _ - .._ .. . .. .. . .. . _ ... .. a - • . Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. significant reduction in multiple cycles[71].These data do suggest that e[CO2]may enhance fruit flavour and fruit yield at the expense of nutritional value. Impact of elevated [CO2]on yield and quality of fruiting trees Sweet clonal cherry(Prunus avium L.)plants were grown for 19 months in climate-controlled greenhouses at ambient(1994-358 ppm;1995- 360 ppm)or e[CO2]('700 ppm).Elevated[CO2]treatment increased photosynthesis and dry matter production,leaf(550/0)and stem(61%),after two months at'7oo ppm,however,this initial stimulation is not sustained.Photosynthetic rates were less after 10 months of growth than after 2 months of growth,and only small increases in dry mass are still evident after 10-months,suggesting that sweet cherry acclimates to e[CO2]due to long- term exposure[72].Due to the young nature of plants studied compared with fully grown mature trees(deciduous tree 15-32 m in height and with a trunk up to 1.5 m in circumference[73,74],no information is available to determine the impacts of e[CO2]on fruit yield or quality.In Nashi pear,a CO2-enriched atmosphere of'700 ppm increased fruit weight,diameter and length along with a 22.5%increase in Brix,(a key measure of sweetness for marketable fruit[75]).However,this also resulted in a reduction in fruit firmness demonstrating that improvements in yield can be nullified by negative impacts on fruit quality(Table 3). While these studies are limited in,they do indicate the potential of CO2-enriched growth for improving photosynthesis,increasing yield and quality of tree crops.However,they also suggest that some crops,especially perennial crops,may become acclimated to higher[CO2]and any gains may be lost over time. Does increasing carbon assimilation increase environmental tolerances? The work presented above also suggest that increasing CO2 uptake could have other benefits.It is notable that growth of fruit crops in carbon enriched atmospheres has a similar effect of protecting against environmental stresses,such as drought and elevated temperature,that may become increasingly common due to climate change as plants genetically engineered to increase carbon assimilation.For example,in melon (Cucumis melo),growing plants in e[CO2]has been shown to mitigate yield losses from increased salinity[67],and in sweet pepper,e[CO2]of 80o ppm was sufficient to rescue any significant yield loss of total and marketable fruits from salinity stress(2o mmol L 1 NaCl)[42].It could be hypothesised that increasing CO2 assimilation increases sugar and chlorophyll content triggering salt tolerance.However,it should be noted that these results are not universally translatable.Gray et al.[76]demonstrated in soybean that e[CO2]was insufficient to protect yields from drought conditions triggered by higher temperatures demonstrating that benefits in some crops may not be translatable across all crops of agronomical importance.Furthermore,in tomato plant Zhou et al.[77]showed that plants grown in e[CO2]were more sensitive to combined drought and heat stress;e[CO2]drives gs and transpiration reducing net photosynthesis and therefore productivity,which is concerning given that greenhouses tend to have elevated temperatures compared to the external environment due to the nature of their construction,glass and metal,and therefore e[CO2] in an enclosed system may negatively impact on yields if water supplies are limiting.This demonstrates that irrigation within greenhouse environments is an essential element and adjusting water regimes to maintain productivity and optimise water-use efficiency. It is also important to note that it is the increase in atmospheric[CO2]that causes the increase in air temperature(along with associated stresses) by absorbing energy and preventing it from being radiated out into space(see[78,79]);as such one might view that the cause cannot mitigate its own effects,however,in some crops where both[CO2]and temperature increase simultaneously,yields were maintained compared with data where temperature is increased in the absence of e[CO2]leading to yield loss and these results cannot be ignored,but a better understanding of the impact of cause and effect climate change on crop yields needs to be researched,otherwise,the logic consequences would be further increase amounts of [CO2]in the atmosphere to increase crop tolerance against the effects of ever-increasing temperatures. Interestingly,some parallels do exist between photosynthetically genetically modified crops and increased tolerance to salinity.In Arabidopsis, over-expression of Sedoheptulose-1,7-bisphosphatase(SBPase),which enhances CO2 assimilation rates by increasing the regeneration of the Rubisco substrate RuBP[8o],enhances salt tolerance through increases in sucrose,starch and chlorophyll content were reported[81].This suggests that increasing photosynthetic rates,either through increasing the availability of[CO2]for photosynthesis or increasing the plants'ability to assimilate[CO2]under ambient conditions could have a similar protective effect.It would be interesting to explore if increased carbon assimilation rates,through atmospheric manipulation or genetic modification,can have a positive impact on crop resistance to high salt environments and other abiotic stresses in large field trials or commercial greenhouses.There is currently evidence that over-expressing the Calvin-Benson cycle (CBC)enzyme SBPase can increase tolerance to chilling stress in tomato[82]and the expression of the cyanobacterial CBC bifunctional fructose- 1,6-bisphosphatases/Sedoheptulose-1,7-bisphosphatase enzyme in soybean prevent yield loss under high temperature[83].Köhler et al.[83] concluded that the manipulation of CO2 uptake could mitigate against the effects of global increases in temperature under e[CO2].This may be deemed especially important given the expected impact of global climate change.This suggests that increasing carbon assimilation through manipulation of photosynthesis[84,85]can have similar outputs to improved photosynthesis through growth in an enriched carbon atmosphere and further demonstrates the viability of this approach for improvement of yield and quality in fruiting crops.This must be studied considering the recent work showing that improved carbon assimilation also results in improved nutrient uptake and an increase in NUE[86]. Future opportunities As[CO2]surpasses 55o ppm,Asat will be limited by the rate of RuBP regeneration rather than Rubisco activity suggesting there is scope to improve plant photosynthesis to increase yield in greenhouse environments where CO2 is routinely increased to 1000 ppm or more for short periods of time. These short time-periods are furthermore unpredictable and chaotic given that greenhouses must be vented,due to external environmental PDF conditions,to maintain,as close as possible,optimal growing conditions i.e temperature and humidity inside the growth facility.Furthermore,the Heip(" [CO2]dosing capacity must be economically beneficial,especially given the chaotic nature of CO2 loss to the environment during periods of venting. Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. £loo per tonne to as much as£3000 per tonne[87,88].Therefore,future options that maximize the ability of the crop to take full benefit of the e[CO2],or maintaining higher yields when CO2 costs are unmanageable become more important. Araus et al[89],noted that canopy photosynthesis holds a crucial place in a context of yield gains through photosynthetic improvement,which requires additional factors including the availability and uptake of nutrients,such as nitrogen,irrigation,the transport of photoassimilates and sink-source balance.As such,in addition to improving photosynthetic rates via CO2 supplementation,the improvement of other plant processes such as N uptake,non-foliar photosynthesis,stomatal function,and rubisco(activase)thermotolerance so that crops are better adapted for growth in[CO2]enriched environments such as greenhouses are discussed below(Figure 3).These works will also need to account for changes to the landscape of greenhouse crop cultivation,such as a move to vertical farming,changes in growth medium from soil to substrates such as coir (derived from coconut husks)or rockwool[9o].It is estimated that more than 5o%of strawberry production occurs in substrate rather than soil [91].Coir is often used as it has been shown to retains water more efficiently than soil,so strawberry plants require less frequent watering improving water use efficiency.Coir also has a high level of aeration,which is ideal for strawberries'whose root systems require a lot of oxygen. More recent developments in hydroponics[92]and aeroponics[93],will impact on irrigation,fertiliser regimes and N uptake. Manipulation of electron Elevation in[CO>] tronspon/� Ambient ICOM 420 + ppm ' Manipulation of stomata density Increased nitrogen assimilation Photosynthetic pigment content manipulation As ICO,)surpasses 550 Wm.A.,:is limited by the CalvimBenson Cycle rate of RuBP regeneration enhanced RuBP regeneration i Is increased I biomass and Increased carbon assimaillionyield possible? and biomass and fruit yield Genetic engineering approaches I} observed in trading pops to reprogram plants for growth in elevated(co( Increased Increased fertilizer uptake. I fertilizer Optimisation of fertilisation Figure 3 Effects of elevated[CO2]on yield of fruiting crops and a representation of the potential for the manipulation of plant material for further yield increases.Created with BioRender.com Nitrogen use efficiency(NUE) With regards to fruit quality,this is a complex trait that may not be simply attributed to enhanced carbon assimilation.More research is needed to link increased assimilate,with assimilate distribution and transport,NUE to better understand the sink-source relationship in any given crop, which can vary significantly across varieties and crop types.NUE is determined by yield per unit of available N in the growth medium(i.e often coir in greenhouse grown crops).Plants with higher NUE may allocate N toward both the photosynthetic complexes(i.e N is major component of chlorophyll;total N allocated to Rubisco 18.2±6.2%[94];)and/or toward the development of additional sinks.The second definition of NUE could be described as the efficiency with which N is applied to soils,(through artificial means in greenhouse crops),is taken up by plants and converted to usable products(i.e.biomass,grain yield).This can be manipulated through breeding to identify new varieties with high NUE uptake from selected growing mediums or through engineering nitrogen symbiosis(Figure 3).Recently,scientists reported the engineering nitrogen-fixation into non- legume cereal crops by enabling them to interact with soil bacteria to convert N from the air into ammonia fertiliser[95].These works could firstly reduce the reliance on commercial synthetic fertilisers and secondly provide alternate sources of N that along with improvements to carbon assimilation,foliar or non-foliar,co-contribute to improving photosynthesis and yields in crops(Figure 3). A recent review has identified a number of targets in the literature to improve N uptake,assimilation and remobilisation through genetic manipulation(see[96]for review).One of these,the over-expression of the nitrate transporter(NRT2.3)was shown to increase nitrate concentrations in tomato increasing biomass and fruit weight[97].More recently,the transcription factor DREBIC has been identified as a regulator of NUE by controlling the expression of several important growth-related genes including the rubisco small subunit 3(RBCS3),nitrate transporters(NRT1.1B,NRT2.4),nitrate reductase(NR2)and the flowering regulator(FTL3).Once over-expressed(OE),OsDREB1C increased the abundance of photosynthetic pigments,plants were shown to have about one-third more chloroplasts,38%more rubisco and improved photosynthesis and N uptake.The OE of OsDREB1C resulted in a>40%increase in grain yield in elite rice varieties and an_20%increase in wheat yields,while in Arabidopsis,a significant increase in biomass[98].Many of these identified genes have potential for improving NUE in fruiting crops grown in e[CO2].A recent report of a large grain rice cultivar,Akita 63,having a high yield due to an enlarged sink capacity without and photosynthesis improvement.However,this work demonstrated that source capacity was strongly limiting the yield potential under high N fertilization.These authors suggested that enhancing photosynthesis is an important step to further increase yield of current high-yielding cultivars[99].This work can be extrapolated that engineering NUE and photosynthesis in plants grown at e[CO2]could provide a step-change in yields in greenhouse cultivated crops. Genetic variation in photosynthetic traits in crops and wild relatives PDF Methods of improving these traits including breeding,by exploiting the potential of crop wild relatives as a source of new traits,and/or the genetic manipulation/genome editing of specific traits.There is already evidence that substantial genetic variation exists within wild relatives of fruiting "Bind III e e— e , . I . IIe - I - e a a'a a a a -O•a •a . •a e --e•e• I e• .11 . a- - •, ,- •, , Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. factors[107,108].This work in wheat and rice is promising,demonstrating the potential for breeding new varieties better adapted to changing growth conditions,however it is unclear if such strategies will work in horticultural crops.In the case of tomato,there is considerable variation within the wild and elite varieties to suggest that such breeding strategies could be used to enhanced yield and quality[109,1101.See Sharwood et al [u1]for review(Figure 3). In transgenic rice,overproducing Rubisco,increases the biomass production and yield under high N fertilization in paddy fields suggesting that the development of new rice varieties with both high photosynthesis and large sink capacity is essential[99].Furthermore,genes encoding thermostable variants of Rubisco activase(thermos-Rca)have been identified in wild rice relatives.When over-expressed in domesticated rice, thermos-Rca was sufficient to enhance carbohydrate accumulation and improve yields after periodic exposure to elevated temperatures(+45°C) throughout the vegetative phase[112,113].Thermostable Rca have been identified in Thermophilic cyanobacteria,bacteria that thrive in high- temperature environments,making them a potential source of novel genes for engineering crops for growth at higher temperatures[114]. Improving the thermal tolerance of rubisco activase,either through breeding with wild populations or genetic engineering,could aid greenhouse grown crops better tolerate the elevated temperatures that often occur during the growing season(Figure 3). Genetic engineering of photosynthetic traits in crops Increasing the expression of enzymes and/or proteins involved in the regeneration of RuBP,CO2 transport or chloroplast electron transport have previously been shown to enhance photosynthetic efficiency and increases in yield[84,85,115-117].However,once again,it cannot be ignored that much of this work has focused on non-fruiting crops,such as Arabidopsis,tobacco,wheat and rice,(see Simkin et al.[84]for review),grown in controlled conditions,performed in pots,in soil or in the field with controlled irrigation,which is not typical of global agriculture.Furthermore, work carried out in tomato,over-expression of sedoheptulose-1,7-bisphosphatase,involved in RuBP regeneration,did not report on fruit yield [82].These data indicating that more work is required to understand how these manipulations would impact fruiting crops grown in tightly controlled environments. One potential target for genetic manipulation is the starch synthesis enzyme adenosine diphosphate glucose pyrophosphorylase(AGPase); increasing AGPase activity has potential to increase starch accumulation for growth.Increased accumulation of starch has been shown to have little negative feedback on photosynthesis[118]and increased AGPase activity in the chloroplast would increase the strength of the transient starch pool, which acts as a sink in the chloroplast.Reduced sink capacity does induce negative feedback on photosynthesis and can limit photosynthesis even in favourable conditions(e.g.elevated[CO2])[119],suggesting that increasing the sink may allow for greater CO2 assimilation in supplemented[CO2] growth environments. Although genetic manipulation has the potential to further increase yields in crops grown in enriched[CO2]environments,allowing them to take better advantage of supplemental CO2 increasing net photosynthetic rates and associated yields(Figure 3),it should also be noted that some reports have suggested that increases in yield in genetically enhanced photosynthetic crops are likely not uniquely down to increases in carbon assimilation but a combination of factors;for example improvements in carbon uptake allow for an increase in N assimilation[120].Furthermore,it has also been reported that such increase in yield from enhanced photosynthetic efficiency critically rely on the availability and uptake of water and nutrients(for review see[121,122]),therefore,genetic engineering as an approach alone may be limiting if other aspects of crop cultivation,such as irrigation,planting regimes,fertilisation(i.e NUE)and growth media(i.e soil,coir,rockwool),are not taken into account and co-optimised. Non-foliar photosynthesis Leaves are not the only location within the plant where photosynthesis occurs,with evidence of photosynthesis in petioles and stems[123,124],and fruit[124]that may provide significant and alternative sources of photo-assimilates essential for optimal yield.Assimilation of atmospheric CO2 is dependent on the number and behaviour of stomata,and the stems of many plants have stomata distributed along the epidermis[125,126]and an evaluation of the photosynthetic activity in stems of various plants accounted for up to 4%of the total photosynthetic activity[127].Furthermore, Hu et al.demonstrated the importance of stem photosynthesis to yield in cotton;maintaining the stem in darkness reduced seed weight by 16% [128]showing the stem provides photoassimilates for plant development and growth. As previously noted,many fruiting crops produce green fruit containing all the necessary proteins and enzymes to carry out photosynthesis[127, 129,130]that may provide significant and alternative sources of photoassimilates essential for optimal yield and quality[124].Tomato fruit photosynthesis contributes to net sugar accumulation and growth and previous work concluded that tomato fruit photosynthesis contributes between lo%and 15%of the total fixed carbon of the fruit[127,131][132],.It should be noted that,unlike many crops,cucumber fruit remain green through to maturity,have stomata(suggesting they perform gas exchange to drive photosynthesis),and have a similar surface area to an expanded leaf[13o].It has previously been reported that cucumber fruit had high photosynthetic and respiratory rates[133]and contribute approximately 9.4%of their own carbon requirements[13o].It should be noted that in fruit with stomata,such as cucumber,there are two potential major sources of CO2.Firstly,Rubisco assimilates atmospheric[CO2]through the stomatal pores,leading to the production of sugars via the CBC and secondly, CO2 released by mitochondrial respiration is re-fixed(recycling photosynthesis)[125,134].Whilst this confirms that photosynthesis occurs in fruits,the extent and importance is not clear.In e[CO2],it seems plausible that cucumber fruit photosynthesis may contribute directly to fruit size (and therefore yield by weight)and quality through their ability to directly access carbon in an enriched atmosphere via their stomata(for a review fruit photosynthesis,see[124,135].Therefore,increasing carbon capture by non-foliar tissues has the potential to significantly impact yield and combined with an increase N uptake(i.e.slow release fertilizers[136])to balance the increased carbon uptake,and optimised irrigation regimes has the potential to maximise such yield gains. PDF HeI1(f Conclusions Oxford University Press uses cookies to enhance your experience on our website.By selecting`accept all'you are agreeing to our use of cookies.You can change your cookie setting More information can be found in our Cookie Policy. maximise productivity.An evaluation of fruit quality under these conditions has also been shown to be highly variable between treatments and difference are observed between cultivars with the same treatment suggesting that much more research is required to identify the specific mechanisms behind changes in fruit quality.In the case of soft fruit production in greenhouse environments,it will be important to determine if the quality of fruit harvested early in the season differs from that of fruit harvested later in the season when plants have spent a more significant period of time exposed to e[CO2]growth conditions.Cherry for instance,when grown under prolonged periods of e[CO21,acclimates to prolonged exposure and initial significant gains in yield observed after two months are less detectable after ten months and are not significantly different to control plants grown at a[CO2][72].This may in one respect account for differences in nutritional quality observed in fruit grown in similar conditions in different studies(i.e.fruit harvested at different times in the study)where additional fertilizer treatments aren't provided. Increases in yield associated with e[CO2]controlled environments maybe about more than additional carbon.Controlled environments also allow the regulation of transpiration(e.g.by controlling vapour pressure deficit)and therefore water uptake and the inclusion of additional fertilisation (specifically N).Breeding new varieties adapted to these growth conditions may also be more amenable given the hostility towards genetically modified crops.A recent review noted that new phenomics,genomics,and bioinformatics tools make it possible to harness the untapped potential of crop genetic resources(including wild relatives)to create combinations of traits to enhance yield in high[CO2]controlled environments[137]. Breeding alone may not be sufficient to adapt all varieties,or all crops,to high[CO2]growing environments traditionally used in greenhouses. However,over the last several decades,agricultural research has adopted technologies such as genetic engineering and"genome editing"to improve traits in key crops that could be useful in these circumstances[85,138-140].These include advances in the tools available to carry out this work,including vectors for multiple gene insertion[141-145]and tissue specific promoters[146-150].If the promise of these biotechnology programs is to be realized,it will be necessary to address the public perception of genetic modification and genome editing technologies to gain greater acceptance. Genetic manipulation,may need to go beyond the direct manipulation of carbon assimilation in leaves[84,180],but focus on the manipulating and control of stomatal function[151,152],the manipulation of pigments complexes in ripening fruit[153],enhancement of light capture by the leaves through the manipulation of chlorophyll distribution and form[1541 and importantly look a methodologies for increasing N uptake via transgenic [96]or traditional means(improved fertilization regimes). It should also be noted that the introduction of new growing,hydroponics,aquaponics and aeroponics may require further study,to breed and adapt or engineer plants root architecture for these new growth media.In conclusion,greenhouse cultivation offers the opportunity to manipulate growing atmosphere,lights and VPD for improved yields and we can now look at the opportunities to breed and engineer plants specifically optimised for these conditions. Acknowledgments This research is funded by the Biotechnology and Biological Sciences Research Council(BBSRC)Collaborative Training Partnerships(CTP)for Fruit Crop Research in partnership with NIAB EMR and Reading University.N.H.D was supported by"Realising increased photosynthetic efficiency to increase strawberry yields"(BBSRC,BB/S507192/1)awarded to A.J.S.A.J.S is supported by the Growing Kent and Medway Program,UK;Ref 107139. Author Contributions N.H.D and A.J.S drafted and wrote the manuscript with input from T.L,C.A.R and C.W who also edited the final version. Conflict of interests The authors declare no competing interests. References 1. Ainsworth EA,Rogers A.The response of photosynthesis and stomatal conductance to rising[CO2]:mechanisms and environmental interactions.Plant Cell Environ. 2007;30:258-70. Google Scholar Crossref PubMed WorldCat 2. Kimball BA.Crop responses to elevated CO2 and interactions with H20,N,and temperature.Curr Opin Plant Biol.2016;31:36-43. 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