2022

Permanent URI for this collectionhttps://hdl.handle.net/1807/109467

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Susceptibility of endangered Cornus florida (Eastern flowering dogwood) to the introduced fungal pathogen Discula destructiva (dogwood anthracnose) in the Canadian Carolinian forest: Insights from environmental, ecological, and population genetics assessments
(Canadian Science Publishing, 2022-10-31) Mitchell, Emily; Fleming, Sage; Dorken, Marcel; Freeland, Joanna
Forest fragmentation and introduced pathogens are negatively impacting trees and forests globally, including the Carolinian forest of southern Ontario, Canada. Multiple species-at-risk live in this threatened but biodiverse forest, including the endangered Cornus florida (Eastern flowering dogwood), which is now limited to fragmented woodlots, and has been decimated by the introduced fungal pathogen Discula destructiva (dogwood anthracnose). Ongoing management of C. florida in Canada is challenged by multiple knowledge gaps, two of which we aimed to address in this study. We first evaluated the association between anthracnose disease prevalence and a suite of ecological and environmental variables across 21 sites. Across our sites, larger trees tended to have the highest disease incidence, and trees on shallow slopes had the most crown dieback. We then quantified genetic diversity and gene flow, and found that genetic structure has not been substantially impacted by habitat fragmentation, although dispersal typically covers short distances. However, genetic diversity is relatively low in smaller populations, and in younger trees. Localized dispersal and eroding genetic diversity may limit future adaptation and hence exacerbate population declines. We recommend that managers prioritize plantings in small populations, avoid shallow slopes, and track younger trees to evaluate age-related mortality.
First report of female gametangia in the invasive macroalga starry stonewort (Nitellopsis obtusa) in North America
(Canadian Science Publishing, 2022-10-29) Harrow-Lyle, Tyler J.; Ginn, Brian K.; Kirkwood, Andrea; Melles, Stephanie J.
Nitellopsis obtusa (starry stonewort) is a macroalga in the Characeae family first documented in North America c. 1974. Since initial introduction, N. obtusa clonal populations quickly established in inland lakes as early as 2005. Despite increased N. obtusa monitoring over the last decade, only sterile or male specimens were documented in North America, however; during routine monitoring in Lake Simcoe and Lake Scugog in 2022, we discovered the presence of female gametangia on N. obtusa. In addition, two other Characeae genera had prevalent antheridia and oogonia, co-occurring with oogonia-presenting N. obtusa, which had not been observed previously despite intensive monitoring since 2008. Further studies in North America are required to confirm the proportion of female populations present within invaded regions, as well as to identify plausible causes shifting gametangia development across non-native and native Characeae, especially within the context of climate change. The presence of oogonia on N. obtusa represents a major change to our understanding of this species and its reproductive ecology in North America.
Transcriptome-wide characterization of alternative splicing in five drug-type cultivars of Cannabis sativa
(Canadian Science Publishing, 2022-11-10) Severson, Tonya F.; Adams, Keith L
Cannabis sativa is widely used for fiber, medicinal, and other purposes, and many cultivars exist, yielding varying proportions of cannabinoids and terpenes. There is considerable interest in characterizing genomes and transcriptomes of C. sativa. Alternative splicing (AS) is a fundamental aspect of gene expression that results in multiple types of mRNAs produced by differential splicing. Transcriptome-wide identification of AS events in drug-type cultivars of C. sativa has not been reported. Here we identified AS events using a transcriptome dataset derived from five drug-type cultivars with divergent chemotypes. Intron retention is the most common event type, followed by alternative acceptor, alternative donor, and skipped exons. We also sought to assess conservation of AS events among cultivars. We found 547 events (5%) unique to a single cultivar, 2661 (25%) shared by 2-4 cultivars, and 7569 (70%) common to all five cultivars. Genes with AS events in each set were analyzed for gene ontology enrichment, showing that genes with AS unique to a single cultivar are enriched for molecular functions related to interactions with ATP, and processes involving transport within cells and across membranes. These results provide insights into the conservation and variation of AS events in multiple cultivars of C. sativa.
Transcriptome-wide characterization of alternative splicing in five drug-type cultivars of Cannabis sativa
(Canadian Science Publishing, 2022-11-10) Severson, Tonya F.; Adams, Keith L
Cannabis sativa is widely used for fiber, medicinal, and other purposes, and many cultivars exist, yielding varying proportions of cannabinoids and terpenes. There is considerable interest in characterizing genomes and transcriptomes of C. sativa. Alternative splicing (AS) is a fundamental aspect of gene expression that results in multiple types of mRNAs produced by differential splicing. Transcriptome-wide identification of AS events in drug-type cultivars of C. sativa has not been reported. Here we identified AS events using a transcriptome dataset derived from five drug-type cultivars with divergent chemotypes. Intron retention is the most common event type, followed by alternative acceptor, alternative donor, and skipped exons. We also sought to assess conservation of AS events among cultivars. We found 547 events (5%) unique to a single cultivar, 2661 (25%) shared by 2-4 cultivars, and 7569 (70%) common to all five cultivars. Genes with AS events in each set were analyzed for gene ontology enrichment, showing that genes with AS unique to a single cultivar are enriched for molecular functions related to interactions with ATP, and processes involving transport within cells and across membranes. These results provide insights into the conservation and variation of AS events in multiple cultivars of C. sativa.
First report of female gametangia in the invasive macroalga starry stonewort (Nitellopsis obtusa) in North America
(Canadian Science Publishing, 2022-10-29) Harrow-Lyle, Tyler J.; Ginn, Brian K.; Kirkwood, Andrea; Melles, Stephanie J.
Nitellopsis obtusa (starry stonewort) is a macroalga in the Characeae family first documented in North America c. 1974. Since initial introduction, N. obtusa clonal populations quickly established in inland lakes as early as 2005. Despite increased N. obtusa monitoring over the last decade, only sterile or male specimens were documented in North America, however; during routine monitoring in Lake Simcoe and Lake Scugog in 2022, we discovered the presence of female gametangia on N. obtusa. In addition, two other Characeae genera had prevalent antheridia and oogonia, co-occurring with oogonia-presenting N. obtusa, which had not been observed previously despite intensive monitoring since 2008. Further studies in North America are required to confirm the proportion of female populations present within invaded regions, as well as to identify plausible causes shifting gametangia development across non-native and native Characeae, especially within the context of climate change. The presence of oogonia on N. obtusa represents a major change to our understanding of this species and its reproductive ecology in North America.
Transcriptome-wide characterization of alternative splicing in five drug-type cultivars of Cannabis sativa
(Canadian Science Publishing, 2022-11-10) Severson, Tonya F.; Adams, Keith L
Cannabis sativa is widely used for fiber, medicinal, and other purposes, and many cultivars exist, yielding varying proportions of cannabinoids and terpenes. There is considerable interest in characterizing genomes and transcriptomes of C. sativa. Alternative splicing (AS) is a fundamental aspect of gene expression that results in multiple types of mRNAs produced by differential splicing. Transcriptome-wide identification of AS events in drug-type cultivars of C. sativa has not been reported. Here we identified AS events using a transcriptome dataset derived from five drug-type cultivars with divergent chemotypes. Intron retention is the most common event type, followed by alternative acceptor, alternative donor, and skipped exons. We also sought to assess conservation of AS events among cultivars. We found 547 events (5%) unique to a single cultivar, 2661 (25%) shared by 2-4 cultivars, and 7569 (70%) common to all five cultivars. Genes with AS events in each set were analyzed for gene ontology enrichment, showing that genes with AS unique to a single cultivar are enriched for molecular functions related to interactions with ATP, and processes involving transport within cells and across membranes. These results provide insights into the conservation and variation of AS events in multiple cultivars of C. sativa.
First report of female gametangia in the invasive macroalga starry stonewort (Nitellopsis obtusa) in North America
(Canadian Science Publishing, 2022-10-29) Harrow-Lyle, Tyler J.; Ginn, Brian K.; Kirkwood, Andrea; Melles, Stephanie J.
Nitellopsis obtusa (starry stonewort) is a macroalga in the Characeae family first documented in North America c. 1974. Since initial introduction, N. obtusa clonal populations quickly established in inland lakes as early as 2005. Despite increased N. obtusa monitoring over the last decade, only sterile or male specimens were documented in North America, however; during routine monitoring in Lake Simcoe and Lake Scugog in 2022, we discovered the presence of female gametangia on N. obtusa. In addition, two other Characeae genera had prevalent antheridia and oogonia, co-occurring with oogonia-presenting N. obtusa, which had not been observed previously despite intensive monitoring since 2008. Further studies in North America are required to confirm the proportion of female populations present within invaded regions, as well as to identify plausible causes shifting gametangia development across non-native and native Characeae, especially within the context of climate change. The presence of oogonia on N. obtusa represents a major change to our understanding of this species and its reproductive ecology in North America.
First report of female gametangia in the invasive macroalga starry stonewort (Nitellopsis obtusa) in North America
(Canadian Science Publishing, 2022-10-29) Harrow-Lyle, Tyler J.; Ginn, Brian K.; Kirkwood, Andrea; Melles, Stephanie J.
Nitellopsis obtusa (starry stonewort) is a macroalga in the Characeae family first documented in North America c. 1974. Since initial introduction, N. obtusa clonal populations quickly established in inland lakes as early as 2005. Despite increased N. obtusa monitoring over the last decade, only sterile or male specimens were documented in North America, however; during routine monitoring in Lake Simcoe and Lake Scugog in 2022, we discovered the presence of female gametangia on N. obtusa. In addition, two other Characeae genera had prevalent antheridia and oogonia, co-occurring with oogonia-presenting N. obtusa, which had not been observed previously despite intensive monitoring since 2008. Further studies in North America are required to confirm the proportion of female populations present within invaded regions, as well as to identify plausible causes shifting gametangia development across non-native and native Characeae, especially within the context of climate change. The presence of oogonia on N. obtusa represents a major change to our understanding of this species and its reproductive ecology in North America.
Transcriptome-wide characterization of alternative splicing in five drug-type cultivars of Cannabis sativa
(Canadian Science Publishing, 2022-11-10) Severson, Tonya F.; Adams, Keith L
Cannabis sativa is widely used for fiber, medicinal, and other purposes, and many cultivars exist, yielding varying proportions of cannabinoids and terpenes. There is considerable interest in characterizing genomes and transcriptomes of C. sativa. Alternative splicing (AS) is a fundamental aspect of gene expression that results in multiple types of mRNAs produced by differential splicing. Transcriptome-wide identification of AS events in drug-type cultivars of C. sativa has not been reported. Here we identified AS events using a transcriptome dataset derived from five drug-type cultivars with divergent chemotypes. Intron retention is the most common event type, followed by alternative acceptor, alternative donor, and skipped exons. We also sought to assess conservation of AS events among cultivars. We found 547 events (5%) unique to a single cultivar, 2661 (25%) shared by 2-4 cultivars, and 7569 (70%) common to all five cultivars. Genes with AS events in each set were analyzed for gene ontology enrichment, showing that genes with AS unique to a single cultivar are enriched for molecular functions related to interactions with ATP, and processes involving transport within cells and across membranes. These results provide insights into the conservation and variation of AS events in multiple cultivars of C. sativa.
Peronospora variabilis is associated with downy mildew of Chenopodium berlandieri in Mexico
(Canadian Science Publishing, 2022-07-26) Félix-Gastélum, Rubén; Leyva-Madrigal, Karla Yeriana; Sánchez-Parra, Luis Roberto; Mora-Romero, Guadalupe Arlene
Peronospora variabilis was observed to be consistently associated with downy mildew of pitseed goosefoot (Chenopodium berlandieri). Morphological characteristics of the conidiophores, conidia and oospores of the oomycete corresponded to those of P. variabilis. The morphological identification was complemented by a phylogenetic analysis of the ITS region. To our knowledge, this is the first report of P. variabilis on C. berlandieri in Mexico.
Trichomycetes from freshwater and salt marsh mosquito larvae with a new species in the monotypic genus Legerioides.
(Canadian Science Publishing, 2022-08-15) Strongman, Doug; Wang, Yan
The intestinal tract of herbivorous arthropods provides a microhabitat for a group of protistan and fungal endosymbionts known as trichomycetes. Many hosts are the aquatic immature stages of insects. One understudied group is mosquito larvae with only six trichomycete species reported from the mosquito gut, but Smittium culicis and Zancudomyces culisetae are globally distributed and common in these insects. Mosquitoes are mostly found in standing freshwater habitats but there are species that are found in brackish water pools common in salt marshes. No trichomycetes are reported from salt marsh mosquitoes. A new species, Legerioides culicicola, is described from mosquito larvae from freshwater sources in Nova Scotia and Prince Edward Island, Canada and Smittium gronthidium is reported for the first time in this host. Smittium culicis is reported from salt marsh mosquitoes from collections in the same provinces in Eastern Canada. Information on the prevalence of mosquito trichomycetes is presented for sites in NS and PEI.
Fungal endophytes affecting the health and recovery of Long’s Braya (Braya longii) and Fernald’s Braya (Braya fernaldii), endangered endemic species of Newfoundland, Canada
(Canadian Science Publishing, 2022-08-15) de la Bastide, Paul Y; Finston, Terrie; Hermanutz, Luise; Hintz, W. E.
Long’s (Braya longii) and Fernald’s (B. fernaldii) Braya are endemic to the Great Northern Peninsula of Newfoundland, Canada. These herbaceous perennials are endangered due to habitat loss and degradation, as well as ongoing threats from fungal infections and insect herbivory. While recovery plans are being implemented, the identity of pathogenic fungi and their origins remain largely unknown. This study was initiated to describe the associated fungal community and infection symptoms. Plant tissues from three populations were sampled over five years yielding 326 fungal isolates identified primarily by nucleotide sequence analysis of the ITS-rDNA region. Isolates included 36 taxa, 16 of which were detected once and 5 detected three times or less. In contrast, 12 taxa were detected across multiple sites, sampling intervals and years. These included two species of Alternaria, Boeremia exigua, closely related species of the Didymellaceae, Botrytis cinerea, Fusarium sp., Colletotrichum sp., Stagonosporopsis sp., Pleospora sp. and Truncatella angustata. Some common taxa showed seasonal trends. The isolated fungi included known pathogens of Brassicaceae. The plants in this study frequently showed symptoms indicative of fungal infection and insect herbivory. These findings improve our knowledge of Braya endophyte communities and will inform recovery efforts for these endangered species.
Comparative ecology of chasmogamous and cleistogamous reproduction in Danthonia compressa: effects of floral type and maternal environment
(Canadian Science Publishing, 2022-10-02) Cheplick, Gregory P.
The significance of maternal and progeny environment to chasmogamy (CH) and cleistogamy (CL) in perennial plants is not well known. The perennial grass Danthonia compressa produces CH spikelets capable of outcrossing on terminal panicles in summer followed by selfing axillary CL spikelets in autumn. This study compared CH and CL seed mass, germination, seedling size, tillering, and reproduction in a sunny woodland edge and shaded interior habitat in the Catskill Mountains of New York State. CL seeds were significantly heavier than CH seeds. Numbers and mass of both seed types were greater in the edge. CL seeds showed much greater germination than CH seeds in both environments during spring, 2018; CL seedlings were larger than CH seedlings in the sunnier habitat. Survival was much greater in the edge than interior and higher for CL seedlings. After 27 months, maternal effects were mostly not detected and there was no significant effect of plant type (CH vs CL) on tiller production, final biomass, or reproduction via CH and CL. For this native grass, non-dispersed CL seeds may be more critical to seedling establishment and population persistence due to greater mass and germinability, and better plant survival, relative to smaller, dispersible CH seeds.
Impact of in vitro hormone treatments on the bibenzyl production of Radula complanata
(Canadian Science Publishing, 2022-05-25) Blatt-Janmaat, Kaitlyn; Neumann, Steffen; Schmidt, Florian; Ziegler, Jörg; Peters, Kristian; Qu, Yang
Bibenzyls are a specialized metabolite class found throughout the plant kingdom. One of the most prolific producers of bibenzyls are liverworts, specifically plants of the Radula genera. These plants possess an incredible diversity of bibenzyls, prenylated bibenzyls, and a few (bis)bibenzyls, several of which have medicinal properties, including perrottetinene, an analog of tetrahydrocannabinol from cannabis. To provide insight into the bibenzyls biosynthesis in planta, exogenous phytohormones were applied to in vitro grown Radula complanata and bibenzyl metabolite production was monitored with targeted and untargeted metabolomics. The targeted metabolomic analysis of six prenylated bibenzyls revealed that production of these metabolites was largely reduced when plants were treated with abscisic acid (AA), salicylic acid (SA), 1-naphthaleneacetic acid (NAA) or 6-benzylaminopurine (BAP). The reduction of these metabolites in the BAP and NAA treatment suggests that prenylated bibenzyl production is negatively correlated with vegetative plant growth. The reduction of bibenzyls at low AA and SA concentrations and mild increase at higher AA and SA concentrations suggests their production is regulated by these stress hormones. In addition, six other bibenzyl metabolites were tentatively identified from the untargeted analysis. These results provide insight into the influence of phytohormones on the bioactive bibenzyl content of Radula complanata.
Diversity among rare and common congeneric plant species from the Garry oak and Okanagan shrub-steppe ecosystems in British Columbia: implications for conservation
(Canadian Science Publishing, 2022-08-02) Hersh, Evan; Wheeler, Erica; Costanzo, Brenda; Ramakrishnan, Alisa P.; Miscampbell, Allyson; Ritland, Carol; Whitton, Jeannette; Gorrell, Jamieson C; Harrower, William Laughton
It is often assumed that the northern periphery of species’ ranges are genetically depauperate due in part to founder effects from post-glacial colonization. The majority of federally protected plant species are peripheral in Canada, yet we have little information about their patterns of genetic diversity and structure. In British Columbia, the majority of these protected plant species occur in two threatened habitats: the Garry oak and Okanagan shrub-steppe ecosystems. Using universal non-coding chloroplast DNA markers, we investigated genetic diversity and genetic structure in four rare and common plant species pairs inhabiting these two ecosystems. We found that rare species had lower genetic diversity than their common congeners, and detected contrasting patterns of regional diversity and structure based on ecosystem. Species from the Garry oak ecosystem showed lower genetic diversity in the northern deglaciated region and significant differentiation between regions, likely due to limited dispersal between Vancouver Island and the mainland. Species from the Okanagan shrub-steppe, however, tended to have uniform diversity across their range and lack regional structure. This study provides an important first look at the phylogeographic patterns of four rare plant species in British Columbia.
Do larger plants produce more and better seeds and seedlings? Testing the hypothesis in a globose cactus, Wigginsia sessiliflora.
(Canadian Science Publishing, 2022-07-22) Ceballos, Celeste; Ferrero, Maria Cecilia; Aliscioni, Nayla L.; Las Peñas, M. Laura; Gurvich, D. E.
In iteroparous plants, an increase in allocation to reproduction is expected with increasing plant size. The aim of this study was to analyze how plant diameter is related to total produced seed mass, seed number, mean seed mass, percentage and mean germination time (MGT), and seedling size and shape in Wigginsia sessiliflora (Cactaceae), a slow-growing globose species from central Argentina. Plant diameter was measured in a population of n185 individuals, and fruits were collected. We counted all seeds to obtain seed number, and weighed them to get seed mass and total seed mass. Seeds were germinated and data of germination percentage and MGT were collected. We also measured the size (height and width) and shape of seedlings (height/width relationship). Fifty four percent of the plants did not produce fruits. Plant diameter was unimodally related to fruit number, total seed mass, and seed number, i.e. intermediate-sized plants presented the highest values. Height and shape of seedlings were positively related to plant diameter. Surprisingly, plant diameter was related in a unimodal way to reproductive outputs. As plants grow, surface-volume ratio decreases, meaning that respiratory losses would be higher than the increase in photosynthetic capacity; therefore, fewer resources may be available for reproduction.
The role of environmental stressors on reproduction, seed morphology, and germination: A case-study of Northern White Cedar, Thuja occidentalis L.
(Canadian Science Publishing, 2022-07-15) Johnson, Sara Ann; Janssen, Eric; Glass, Nicholas; Dickerson, Patricia; Whelan, Christopher J.; Molano-Flores, Brenda
Changes in resource allocation from parent to offspring can have effects on the dispersal, recruitment, and survival of progeny, with impacts sometimes extending across generations. In plants, environmental stressors not only impact the growth of mature individuals but can also alter the provisioning of resources to developing seeds, further extending to embryo development and reproduction. In this study, we compare seed biomass, morphology, seed set, and germination among two natural populations and three additional seed sources of Thuja occidentalis (Northern White Cedar), a fen-dependent species in the Chicago IL region. Chicago Junior School and Trout Park were once contiguous populations, now bisected by the I-90 toll road, and are heavily impacted by human disturbance and external pollutants. Variation was observed in seed morphology, cone biomass, and seed set among study sites with Trout Park trees producing fewer seeds and smaller cones than any other site in our study. Despite morphological differences, germination was low overall except for commercial seed. Differences among the two neighboring sites implicate that Trout Park is disproportionately affected by environmental factors (e.g., road salts and altered hydrology) compared to Chicago Junior School, potentially impacting the growth and recruitment of Thuja occidentalis in these urban populations.
Spatial separation of mycobionts in the giant, differentiated root system of Cyrtosia septentrionalis, a fully myco-heterotrophic orchid
(Canadian Science Publishing, 2022-06-26) Umata, Hidetaka; Ota, Yuko; Gale, Stephan William; Chuman, Sadaharu; Nishi, Munetaka; Ashihara, Seiichi; Yagi, Fumio
The rare achlorophyllous orchid Cyrtosia septentrionalis is thought to switch mycobiont from a Physisporinus species (‘TK-10’) to an Armillaria species as it develops. We conducted morphological, chemical and molecular analyses of the root system of three adult plants growing sympatrically in Japan to assess reliance on different mycorrhizae. The root systems were found to comprise several long primary roots that radiate outwards from the emergent stem and many shorter lateral ones. Long roots branched most vigorously where they encountered A. gallica in the soil, and the resulting lateral roots contained the highest density of intra-radical rhizomorphs. One plant was associated with TK-10 and the other two with A. gallica, but only the apical portion of long roots and a few lateral roots were colonized. Mycobionts isolated from the base of long roots proved incapable of germinating C. septentrionalis seeds. Whereas glucose was the dominant monosaccharide in non-mycorrhizal roots, galactose was in mycorrhizal ones. Lateral roots are specialized in mycorrhization and their velamen and mucilage may play a key role in preventing desiccation. We hypothesize that C. septentrionalis can associate with either fungus from germination to adulthood, but that detection of A. gallica in the soil can prompt replacement of TK-10.
New insight into the roles of lipid transport protein and seed storage albumin gene families involved in oil and protein accumulation in rapeseed (Brassica napus L.)
(Canadian Science Publishing, 2022-01-16) Chen, Chuanhong; Chen, Lunlin; Li, Shuyu; Zhai, Wen; Li, Dongwen; Chen, Jun
In rapeseed, lipid transfer protein (LTP) and seed storage albumin (SSA) gene families are involved in lipid and protein metabolism. Understanding the mechanism of oil biosynthesis have great value to increase oil production. The LTPs and SSA gene families were identified of Brassicaceae. The LTPs were divided into 6 groups according their sequences, and LTPs of all five species in Brassicaceae also had the same pattern. The genes at a higher expression level were evenly distributed across 5 groups except group ii. The gene groups in these two gene families showed different expression patterns, with most of the LTP genes expressed at a higher level at 25 days after flowering (DAF), but SSA genes were highly expressed at 40 DAF stage. Gene ontology (GO) enrichment analysis of the regulatory genes for LTPs also was performed, including response to biotic stimulus (GO:0009607), generation of precursor metabolites and energy (GO:0006091), electron transport chain (GO:0022900), oxidative phosphorylation (GO:0006119), ATP metabolic process (GO:0046034) and phosphorylation (GO:0016310). These results may facilitate further research to understand the expression patterns and regulatory mechanism of LTPs and SSAs.
Population-level drought responses in northern oak fern (Gymnocarpium dryopteris)
(Canadian Science Publishing, 2022-04-27) Winkel, A.; Wood, Lisa J
Ferns serve as an invaluable indicator species for soil moisture and distribution patterns of other plant groups. Northern oak fern (Gymnocarpium dryopteris), a common fern in the interior of British Columbia, Canada, has received little research attention despite its prevalent use in the forest resource sector as an indicator species for moisture. To understand the impact of water deficits on oak fern, we sampled in two different ecoregions in interior British Columbia and assessed acclimation to drought, by measuring the size and abundance of stomata in each region. We then subjected the ferns to a period of drought, and recorded the response to stress by measuring the photosynthetic efficiency of the oak fern populations over time. Differences existed in the morphology of oak fern populations; ferns from the wetter ecoregion possessed more, larger stomata then those from the drier ecoregion. Oak ferns from both regional populations were significantly reliant on moisture present in the soil and air to maintain photosynthetic efficiency, and

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