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Figure 4 illustrates the length of fertile internodes 20–40 within the various treatments of: FC, FN, F45, T45, N45, and B90. The FC, FN, and F45 treatments grew lengthier internodes from node 20–40 than the T45, N45, and B45 treatments (Fig. 4). Internode width, however, was greater in T45, N45, and B90 as compared to the undisturbed FC, FN, and F45 treatments (Table 1). The T45 and N45 treatments had a 27.9% and 26.6% reduction in internode elongation compared to the FC, FN, and F45 treatments. Of the treatments, B90 had the shortest internodes and widest internode thickness between node 20–40 (Tables 1, 2). Due to the shorter internode lengths in the mechanically affected treatments, the density of nodes per unit area was ~ 25% greater in T45 and N45 from node 20–40 and an additional 28% shorter in B90. In other words, B90 internodes were ~ 54% shorter between nodes 20–40 as compared to the untouched treatments and had the densest node concentration (cf Fig. 4 and Table 2). Both touched and bent bines were significantly reduced in elongation (Table 1; P  12–25. Thus, amassing many fertile nodes per vertical distance within a high sidewall greenhouse (e.g. ≥ 6 m) would be one viable means to increase the yield potential of hop in controlled environment production as long as plant resources did not become limiting. What’s more the 15.25 cm rise over run staircase created by the B90 internode bending treatment would allow for approximately double the bine length from the container to the top of a high sidewall greenhouse as compared to a vertically trellised bine (an additional direct step toward increasing node quantity per unit vertical production area). Secondly, the time and resource investment in overcoming the hop cultivar specific 11–24 infertile juvenile phase adds approximately three weeks to a single hop crop cycle e.g.11,36. Thus, it would be more time and space efficient to grow fewer crop cycles per annum that contain larger amounts of fertile nodes within a cycle as compared to additional cycles that contain the unfertile juvenile phase.In conclusion, repeated touch and/or bine bending within the active elongation zone of hop bines resulted in shortened internode length with higher cone production per given area. Mechanical stimuli did not reduce cone yield or flower quality. The results demonstrate that successive local internode strain can aid the control of internode elongation. Moreover, the study provides evidence that thigmomorphogenic cues can be used as a management tool to increase bine compactness and increase node density per unit area. This finding is especially important for growth control when production space is limiting and/or of high-value (e.g. greenhouse production)1. Hence, mechanical perturbation was an effective non-chemical means to control hop internode length. Nonetheless, models aimed at predicting internode length of hop bines in response to strain should still take into account a cultivar parameter. The results are practical on a commercial scale because the methods of touch and bending used in this study are easy to apply with minimal investment in labor, have a short time interval of application (approximately 5–10 s−1 per bine per 24 h), and the application duration is relatively short ~ 30 days out of the 90–120 day crop cycle, making this a practical endeavor when one considers that high value vine crops are already repeatedly handled by humans throughout their production cycle (e.g. viticulture grape and controlled environment cucumber production). More

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Plant materialsIn April 2019, 2-year-old Chinese fir seedlings were collected from the Yalin Center of the Chinese Academy of Forestry in good condition and free from pests and diseases. (The use of Chinese fir seedlings in the experiment complies with national regulations).MediumPikovskava (PVK) solid medium: glucose 10 g, Ca3(PO4) 25 g, CaCO35g, (NH4)2SO40.5 g, NaCl 0.2 g, MgSO4 7H2O 0.1 g, KCl 0.1 g, MnSO4 0.002 g, FeSO4 7H2O 0.002 g, agar 18 g, Distilled water 1000 mL, pH 7.0.PVK liquid medium: PVK solid medium without agar.Luria-Bertan (LB) medium: tryptone 10 g, yeast extract powder 5 g, NaCl 10 g, agar 18 g, distilled water 1000 mL, pH7.0.LB liquid medium:LB solid medium without agar.Isolation and purification of endophytes from Chinese fir seedlingsThe roots, stems, and leaves of the selected Chinese fir seedlings were washed away with running water to remove the surface soil, and then washed with running water for 24 h to 36 h, and the surface moisture was absorbed with sterile filter paper. Weigh 1 g of roots, stems and leaves in a petri dish, and then carry out surface disinfection in a sterile operating table with 75% alcohol (C2H5OH) for 30 s, 5% sodium hypochlorite (NaClO) 10 min, wash the sterile water 7 times, and use sterile filter paper to absorb the water. The material after surface sterilization is cut into 2 mm × 2 mm with sterile surgical scissors, placed in a sterilized mortar and grated with a small amount of sterile quartz sand and ground into a homogenate, then diluted with sterile water to 10–1, 10–2 and 10–3, pipette to draw 150 µL of sample grinding fluid, spread on the medium, set the sterile water of the last rinse of Chinese fir tissue as a blank control, incubate with other plates under the same conditions, and verify whether the surface disinfection. Cultivate in a 28 ℃ incubator according to the characteristics of the colony phenotype, and use the streak separation method to further purify and isolate the strains until the isolation of the colony morphology is uniform for each isolate.Note: 75% C2H5OH: 75 mL absolute ethanol + 25 mL sterile deionized water; 5% NaClO: sodium hypochlorite solution with 10% available chlorine: sterile deionized water = 1:1.Characterization of PGP traitsDetermination of phosphorus solubilizing abilityThe strain was inoculated into PVK liquid medium and cultured at 28 ℃ for 180 days/min on a reciprocating shaker for 7 days, and then the pH value of the medium was measured. The culture solution was centrifuged at 8000 rpm for 15 min to remove bacterial cells. Take the supernatant and use the molybdenum antimony scandium colorimetric method23 to determine the soluble phosphorus content in the culture broth.Determination of nitrogenase activityAn aliquot of 200 µL fresh culture was inoculated to 20 mL of nutrient broth and incubated overnight at 30℃. Bacterial growth was collected by centrifugation and was washed twice using sterile water, and resuspended by liquid limited nitrogen culture medium (OD600 = 0.2). The 3 mL suspension was transferred to a 25 mL sterilized serum vial and 2.4 mL acetylene gas (99.9999%) was driven into the serum bottle, and then incubated at 30 °C for 12 h. The ethylene content and the protein of bacterial suspension were determined as You et al.24.1-Aminocyclopropane-1-carboxylate (ACC) deaminase activity determinationACC deaminase activity was determined by the method of Glick et al.25 using N-free medium (Nfb)26 for bacteria and minimal medium (MM)27 for actinomycetes containing 0.3 m mol L−1 ACC (Sigma, USA) as a sole nitrogen source. MM with 0.1% (w/v) NH4(SO4)2 was used as a positive control and cultivation without ACC was used as a negative control. After incubation at 28 ℃ for 7 days for non-actinomycete bacteria and 14 days for actinomycetes, colony growth on Nfb or MM with addition of ACC indicated ACC deaminase activity.Indole-3-acetic acid (IAA) productionIAA production was measured by colorimetric assay27. Bacterial isolates were cultured for 3 days in TY broth (without L-tryptophan or supplemented with 500 μg/mL of l-tryptophan) in the dark at 28 °C. Cells were removed from the culture medium by centrifugation at 13,000×g for 10 min; then, 1 mL of the supernatant was mixed vigorously with 2 mL of Salkowski’s reagent (4.5 g of FeCl3 per L in 10.8 M H2SO4). Samples were incubated at room temperature for 30 min and the IAA production was estimated from the optical density at 600 nm (OD600) by comparison with a standard curve prepared from known concentrations of IAA.Siderophore productionSiderophore production was examined by using chrome azurol S (CAS) agar28. Isolate was inoculated onto CAS agar, cultured at 28 °C for 2 days, and the positive strain was indicated by an orange halo around the bacterial colony. Determine the ratio (D/d) of orange aperture diameter (D) to colony diameter (d) to determine the iron-producing carrier capacity of the strain.Physiological and biochemical tests of phosphate-solubilizing bacteriaThe conventional physiological and biochemical identification of PSB is carried out according to the methods in the “Common Bacterial System Identification Manual”, which mainly includes Gram stain, glucose hydrolysis test, lactose hydrolysis test, methyl red test, Voges-Proskauer (VP) test, hydrogen sulfide production test, gelatin liquefaction Test, citrate utilization test, malonate utilization test, denitrification test.16 SrRNA gene sequencingTaking the screened multifunctional PSB as the object, the bacterial genomic DNA extraction kit of Beijing Bomed Biotechnology Co., Ltd. was used to extract the DNA of the strain, using the DNA as a template, and using the bacterial universal primer 27F (5′-AGAGTTTGATCCTGGCTCAG-3′) and 1492R (5′-GGTTACCTTGTTACGACTT-3′) PCR amplification, the amplification system is as follows: DNA template 1 uL, primer 27F 0.5 µL, 1492R 0.5 µL, 2 × TaqMix 12.5 µL, ddH2O10.5 µL. The PCR procedure is as follows: 93℃ for 3 min, 93℃ for 30 s, 56 ℃ for 30 s, 72℃ for 2 min, 32 cycles; 72 ℃ for 7 min. The amplified products were sequenced bidirectionally by BGI. After splicing the measured 16SrDNA sequences in ContigExpress, search in GenBank, EzTaxon, BIGSdb databases respectively, select the model strains with high homology. The phylogenetic tree was constructed by the neighbor-joining method using MEGA version 7.0 with the Kimura 2-parameter model29, the robustness of the tree was evaluated by performing bootstrap analyses based on 1000 replications30.Evaluation of plant growth promotion by individual inoculationPreparation of inoculumThe single colonies were picked out and incubated in LB broth at 180 rpm at 28 ℃ for 12 h. The above solution was inserted into 200 mL of LB broth at 1% inoculation, incubated at 180 rpm at 28℃ for 48 h and the cell pellet was resuspended in sterile distilled water and made up to a final concentration of 3 × 108 CFU/mL.Experimental seedlings and soilChinese fir seedlings were provided by the Experimental Center of Subtropical Forestry, Chinese Academy of Forestry (117° 67′ E, 27° 82′ N), Jiangxi Province, China. The use of Chinese fir seedlings in the experiment complies with national regulations. Five months old seedlings with vigorous and apparently disease and pest free were used. The height and root collar diameters of seedlings were 8.7 cm and 1.36 mm, respectively. The seedling container was made of non-woven fabric, the specification was 4.5 cm × 8.0 cm (Diameter × Height). The soil was consisted of nursery medium and loess at a ratio of 9:1, was thoroughly mixed and homogenized with 3 kg slow-release fertilizer per cubic. The slow-release fertilizer is produced by American Abbes (180 g kg−1 total N, 80 g kg−1 available P, and 80 g kg−1 total K, the fertilizer effect period is 9 months). The soil exhibited the following properties: 6.34 g kg−1 total N, 0.80 g kg−1 total P, 2.50 g kg−1 total K, and a pH-value of 6.00.Test designThe SSP2, JRP22 and HRP2, which were confirmed to have the characteristics of promoting plant growth, so pot experiments were conducted. To determine the effectiveness of phosphorus-solubilizing bacteria in plant growth of Chinese fir, a pot culture experiment was conducted between August and November 2019 in an open-sided greenhouse in Experimental Center of Subtropical Forestry, Chinese Academy of Forestry, Jiangxi Province, China. The experiment was carried out in three-factor orthogonal design with five replications for each treatment. The orthogonal experimental design of experiment is provided in Table 1, and strain, dilution ratio, inoculation method contained 3 levels. The pots with water was used as control (CK). For the irrigation of the rhizosphere (IR) treatments, 30 mL of diluted inoculum was added to the soil in the vicinity of the roots of Chinese fir. For the foliar spray (FS) treatments, 30 mL diluted bacterial cell suspension was inoculated in the leaves of Chinese fir by using a syringe. For the rhizosphere + foliar spray (IS) treatments, 15 mL of diluted inoculum was inoculated in the rhizosphere of seedlings, 15 mL was added to the leaves of seedlings by foliar spray. Each treatment contained sixteen seedlings for a total of nine treatments. A total of 3 inoculations were given in the middle of each month. The plant height and stem diameter were recorded before the first inoculation. The plants were harvested after 90 days (16 plantlets/replicate/treatment, i.e., a total of 80 plantlets per treatment) and the root biomass, stem biomass, leaf biomass, plant height and stem diameter were measured.Table 1 L9(34) Orthogonal design of experiment.Full size tableDetermination of growth indicatorsDuring the test, before each inoculation, the height of the seedlings was measured with a ruler and the ground diameter of the seedlings was measured with a vernier caliper. After the experiment, 10 plants of Chinese fir seedlings in average growth were randomly selected from each treatment, a total of 30 plants were washed with clean water to remove surface impurities, the filter paper was dried and the roots, stems, and leaves were put into paper bags respectively at 105 °C. After being degraded for 0.5 h, dried at 70 °C to a constant weight, weighed and recorded the biomass of each part.Determination of leaf and soil nutrient contentTotal N content of leaf and soil was measured by a 2300 Kjeltec Analyzer Unit (FOSS, Höganäs, Sweden). Total P and total K, total Mg and total Fe of leaf, soil TP, TK, AP and AK were extracted according to literature31 and were determined by ICP (Kleve, Germany).Determination of soil enzyme activitiesActivities of the soil urease, cellulase, sucrase, dehydrogenase and acid phosphatase were determined by spectrophotometry. Firstly, 0.05 g of soil was added to 450 mL of phosphate buffer solution (PBS, 0.1 mol L−1, pH 7.4). Then the solution was mixed by shaking, and centrifuged at 2000 rpm at 4 ℃ for 10 min and supernatant was collected with a new centrifugal tube. The supernatant and reagents were added according to the kit instructions (Shanghai Enzyme-linked Biotechnology Co., Ltd., Shanghai, China). Absorbance at 450 nm was measured on a SpectraMax Paradigm Multi-Mode detection platform (Molecular devices, San Jose, CA, USA).Data processing and analysisAll statistical analyses were performed using SPSS24. Data are presented in terms of means (± SE; standard error). Statistical differences were tested by one-factor ANOVA to evaluate the differences in the nutrient content of soil and plant growth status. In MEGA7.0, the Neighbor-Joining method was used to construct the phylogenetic tree, and the Bootstrap value was 1000. More

Vegetation history of the Acıgöl areaOur palynological analyses of 72 regularly spaced samples show a diversified vegetal landscape alternately wooded and open, in response to orbitally driven climatic cyclicity. However, arboreal pollen values remain almost constantly below 50% of the Pollen Sum (PS) (average 27.5%, median 22.8%), which corresponds to an overall open landscape (Fig. 3). Among herbaceous plants, the dominant taxa are steppics such as Artemisia, heliophilous and halophilous taxa including Calystegia, several Compositae, Convolvulus, Linum, Plantago ssp., Poaceae and Chenopodiaceae that could develop on the saline shores of Acıgöl lake during evaporitic periods. Forests are composed of a mixture of conifers, Mediterranean Pinus, Abies, Cedrus, Cupressaceae and Picea, associated with broadleaved trees dominated by Mediterranean oaks, i.e. deciduous and evergreen Quercus, with some Olea. Riverine trees such as Alnus, Salix, Populus, Tamarix, Juglans and Platanus have also been identified. Few Tertiary or megathermic relictual taxa (Carya, Liquidambar, Parrotia, Pterocarya fraxinifolia, Taxodiaceae, Tsuga, Zelkova) were identified so far in the pollen assemblages, mostly before 2.2 Ma, due to climatic cooling17,18 since the end of Tertiary which led to a decline in global biodiversity19,20.Figure 3Simplified pollen and NPP diagram in percentages of Acıgöl, core 3, based on the age model of Demory et al. [1]. Equidistant scale. Values are in percentages calculated on a pollen sum without Non-Pollen Palynomorphs (NPP), Ferns, Bryophytes and Algae. The beige rectangle corresponds to the date of the presence of Homo erectus at Kocabaş (Lebatard et al. [4]).Full size imageThe vast freshwater stretch of Acıgöl, located in a predominantly arid limestone hills environment, seems to have been a crucial resource for the mammalian fauna, which probably concentrated around the site in search of water and pastures. Indeed, low percentages of arboreal pollen imply that the landscape remained open throughout the sequence and suggest a marked grazing pressure by herbivores in addition to climatic factors21,22,23.Coprophilous fungi spores, cereals and other ancestors of cultivated plantsCoprophilous fungi spores are excellent indicators of herbivorous mega-mammal herds since they grow exclusively on dung deposited by these animals24. At Acıgöl, a wide variety of coprophilous fungi spores has been identified throughout the pollen record including: Sporormiella sp., Podospora sp., Delitschia sp., Sordaria sp. and Valsaria variospora (Figs. 3, 4). They provide evidence for a continuous presence of large herbivorous mammals around the lake throughout Quaternary.Figure 4Coprophilous fungi spores of Acıgöl, core 3. Equidistant scale. Age model is from Demory et al. [1]. In red: coprophilous fungi taxa..Full size imagePollens of Poaceae, such as Secale (rye) and Cerealia-type, have been identified throughout the sequence (Figs. 3, 5). Unexpectedly, they present the same morphological characteristics as that of modern cereal grains25,26, namely an average size of ≥ 40 µm and a large pore + annulus (≥ 8 µm). As by definition cereals are cultivated plants, we will call the corresponding plants “proto-cereals” to highlight that their pollen are identical to those of cereals. This resemblance can be seen clearly in Fig. 5, where we have brought together fossil cereals from Acιgöl (Fig. 5, photos 1–7), from Roman time (Fig. 5, photo 8), not modified by modern agricultural practices, and from the current wheat field of the Lauragais agricultural plain, Gardouch, France (Fig. 5, photo 9). Cerealia-type frequencies reach a maximum of 9% of the PS around 2.2 Ma and can be as abundant as wild Poaceae pollen (Fig. 3). The Cerealia/Poaceae ratio shows that 24.66% of all Poaceae are proto-cereals from 2.0 to 2.3 Ma (Supplementary Table 1). Such high proto-cereal rates are almost never reached in pollen records, even in recent periods and in the presence of agriculture, because of the very low pollen dispersal capacity of cereals27. A lowering of frequencies down to 2–4% range is recorded in younger periods (Fig. 3), as well as a step like decrease of the Cerealia/Poaceae ratio (Fig. 6). This change may be related to the Middle-Pleistocene Transition (MPT) cooling and to the mega-mammal fauna change from a Villafranchian to a Galerian type28. MPT and faunal changes occurred around 0.9–1.0 Ma, while a decrease in our proto-cereal starts around 1.5 Ma, however signs of cooling and amplified climatic cycles predate the MPT28.Figure 5Pollen grain of Cerealia and Triticum sp. from Acıgöl (ACI), core 3 (photos 1–7), the Roman site of La Verrerie, Arles, France (photo 8) and Gardouch, France, current wheat field (photo 9). Photographies with a photonic (photo 1 – 4 and 8) and a confocal microscope (photos 5-7 and 9). 1) sample ACI 239 m, age: 0.871 Ma. 2) sample ACI 435.50 m, age: 1.709 Ma. 3) sample ACI 532.44 m, age: 2.122 Ma. 4) sample ACI 509.50 m, age: 2.026 Ma. 5) sample ACI 552.57 m, age 2.206. 6) sample ACI 552.57 m, age: 2.206 Ma. 7) sample ACI 429.50 m, age: 1.681 Ma. 8) sample La Verrerie 1455, age: 50-70 BC (Roman). 9) current pollen of Triticum sp., age: 2000 AD. L: maximal length (µm).Full size imageFigure 6Cerealia/Poaceae ratio in %, % cultivated tree ancestors and % Olea of Acıgöl, core 3.Full size imageThe histogram of wild Poaceae and proto-cereal pollen size (Fig. 7a) shows that there are a number of pollen populations modes around 30, 37.5, 45–50, supporting the idea that the larger grain sizes cannot be interpreted as a tail of ‘anomalous’ wild Poaceae pollen. Moreover, comparison with the present-day pollen rain recorded in moss pollsters, sampled around the lake of Acıgöl (Fig. 7b and Supplementary Table 2), show that the large pollen size mode (≥ 40 µm) is nowadays nearly absent (0–0.97% of the PS, Cerealia/Poaceae ratio of 4.52%, Supplementary Tables 3 and 4), even in biotopes with wild Poaceae considered to be ancestors of cereals (Aegilops, sample 2a, cereal rate: 0.97% of the PS) or with cereals such as Hordeum (sample 3a, b and 4, cereal rate 0.31, 0.00, 0.33 of the PS respectively, Supplementary Tables 2 and 3).Figure 7a) Pollen size of wild Poaceae and proto-cereal of Acıgöl, core 3. The measurements were made on the 10 samples with the highest cereal pollen content. A total of 991 grains of pollen were measured. b) Current pollen rain at the Acıgöl lake and surroundings. 8 moss samples were collected and 354 measurements of the longest axis of the wild Poaceae and cereal pollen grain were made.Full size imageOur interpretation is that proto-cereals recorded throughout the Acıgöl sequence derive from wild Poaceae. Their emergence and predominance may have been favoured by the impact of large herbivore herds attracted to Acıgöl lake shores, and through genetic drift. Through the process of trampling, nitrogen enrichment of soils and browsing, large mammal herds could have altered the genotype of proto-cereals naturally present in Acıgöl and thus, favoured the emergence of modern cereals. For genetic reasons, the descendants of these proto-cereals are not represented today among cultivated Poaceae because domestication bottlenecks eliminate genetic variation29.Is there a relationship between the size of proto-cereal pollen and climate? To our knowledge, the genetic literature does not show any relationship between the increase in pollen size and temperature. However, there does seem to be a relationship with atmospheric drought30,31 which is said to have favoured the appearance of polyploidy in certain species of Poaceae. It cannot be excluded that climate has had an influence on the proto-cereal genome, but only the interaction between herds of large herbivores and proto-cereal steppes can explain why proto-cereal pollen has never been found in such abundance elsewhere in Pleistocene pollen records.The ancestors of cultivated trees (Olea sp., Juglans sp., Castanea sp., Corylus sp., Prunus t.), typical of the modern Mediterranean agriculture, are also present in the Acıgöl sequence (Fig. 3 and Supplementary Table 5). Their amount increases after 1.5 Ma, mainly due to Olea (Fig. 6). Other potentially edible plants such as Ephedra, Hippophae, all the Compositae and the Fagaceae have been identified in the pollen assemblages. They correspond to 54.4% of plants identified in the pollen assemblages. Among these plants, there are 72% grasses and 28% trees and, among edible organs, 51% are vegetables and 20% are seeds (Supplementary Fig. 1a,b). These results testify to the potential wealth of accessible food resources that human and animal populations could feed on. Interestingly, studies carried out in Spain on the present-day consumption of wild plants lead to results close to those obtained at Acıgöl, with 87% grasses and 13% trees32.In recent years, new biological and archaeological data obtained from sites with human occupation have improved our knowledge of the beginnings of agriculture and the modalities of its implementation. In the Levant, the Ohalo II site highlights the presence of proto-cereal seeds, and flint tools to harvest, as early as 23,000 years before the present33. Further north, on the archaeological site of Gesher Benot Ya’aqov, proto-cereal seeds (oats, Avena) as well as pollen from cereals and trees currently cultivated, were identified over a period ranging from 750,000 to 820,000 years34,35. Moreover, recent genetic data indicate that the emergence of agriculture did not occur at a single location at the onset of the Neolithic (e.g. the “Fertile Crescent” hypothesis) but is, on the contrary, an evolutionary and multi-regional long-term phenomenon36,37,38. Alternatively, or simultaneously, are the hominins also partly responsible by having developed episodes of a form of transitory “proto-agriculture”? We already know that this domestication process was discontinuous with shutdown and restart phases37,39. Acheulean lithic tools, characterised by symmetrically shaped bifaces, testify to the rather advanced cognitive capacities of early Pleistocene populations that may have visited the lakeshore of Acıgöl5. Hominin populations may also have benefited from this opportunity to diversify their food regime with easily harvested and nutrient-rich wild plants (Supplementary Table 5), as it is the case today for hunter-gatherer populations in Africa and elsewhere in the world. More

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