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PubMed2003

Activated sludge as inoculum for ready biodegradability testing: effect of source.

Results of ready biodegradability tests (RBT) are barely reproducible owing to a well-known lack of definition in inoculum source and quality. In this study, the degree of variability expected when only activated sludges are used as inoculum source was investigated. For this, the characteristics of activated sludges collected in municipal wastewater treatment plants operating at various massic loading rates (MLR; 0.1, 0.5 and 0.9 kgBOD5 kgVSS(-1) d(-1)) were compared. In order to provide suitable cellular densities for RBT, inocula were obtained after settling of activated sludges and analyzed in terms of active and cultivable cell densities, dehydrogenasic activity, BOD5 and a general profile of hydrolytic enzymes. In our analysis, biomass obtained from the High-MLR treatment plant constituted the inoculum having the highest biodegradation potential both with respect to microbial densities and to enzyme activities. This biomass also yielded the fastest biodegradation kinetics in dodecyl benzene sulfonate RBT. An attempt of biomass homogenization of inocula on the basis of cultivable cell density and dehydrogenasic activity gave negative results with this chemical compound. Since, in practice, restriction of activated sludge sources may be difficult, our results emphasize the importance of further studies aimed at homogenization of inoculum quality and quantity.

Biodegradation, Environmental↗

PubMed1993

Penicillin-binding protein expression at different growth stages determines penicillin efficacy in vitro and in vivo: an explanation for the inoculum effect.

Mechanisms to explain the "inoculum effect" have not been elucidated in gram-positive infections. A mouse model of group A streptococcal myositis was used to compare the efficacies of two beta-lactams, penicillin and ceftriaxone, and a protein synthesis inhibitor, clindamycin, at three different inoculum sizes. beta-lactams were more susceptible to inoculum effects than was clindamycin both in vivo and in vitro (P < .05). The large inocula were hypothesized to reach stationary phase of growth sooner than smaller inocula both in vitro and in vivo. The penicillin-binding protein (PBP) patterns from membrane proteins isolated from mid-log-phase and stationary-phase cultures of Streptococcus pyogenes were compared. Binding of radiolabeled penicillin by all PBPs was decreased in stationary cells; however, PBPs 1 and 4 were undetectable at 36 h. Thus, the loss of certain PBPs during stationary-phase growth in vitro may be responsible for the inoculum effect observed in vivo and may account for the failure of penicillin in both experimental and human cases of severe streptococcal infection.

Animals↗

PubMed1987

The inoculum effect with gram-negative bacteria in vitro and in vivo.

The in-vitro MICs and MBCs of ciprofloxacin, cefoperazone, ceftazidime, ceftriaxone, gentamicin and imipenem were measured for five strains each of Escherichia coli, Klebsiella pneumoniae and Pseudomonas aeruginosa with standard and high inocula. The MBC was raised more than the MIC at the higher inoculum for all the drugs except ciprofloxacin. However, the MBC of imipenem against P. aeruginosa was increased relatively little at the higher inoculum in contrast to the findings with other beta-lactam drugs. In a rabbit model of endophthalmitis both cefoperazone and imipenem were markedly more bactericidal against a high inoculum of K. pneumoniae than would have been predicted from the results of in-vitro time kill curve studies. We conclude that the therapeutic relevance of the inoculum effect remains to be proved.

Animals↗

PubMed2001

Effect of inoculum form on in vitro antifungal susceptibilities of Aspergillus spp.

The effect of inoculum form, i.e. ungerminated conidia, germinated conidia and hyphae on the in vitro antifungal susceptibility of 12 strains belonging to Aspergillus spp. was compared using a broth microdilution method. The isolates were tested three times on different days against amphotericin B, itraconazole and ketoconazole. There were no significant differences between MICs obtained with the three types of inoculum (P > 0.05) for any antifungal tested. The degree of reproducibility of the tests was high (>or=75%) for all antifungals with each type of inoculum, except for itraconazole (58.3%) when the inoculum was prepared with ungerminated conidia.

Amphotericin B↗

PubMed2004

Effect of inoculum density on susceptibility of Plesiomonas shigelloides to cephalosporins.

OBJECTIVES: Resistance of Plesiomonas shigelloides to cephalosporins at higher cell densities has been reported. We investigated whether these inoculum effects are due to the production of beta-lactamases. METHODS: beta-Lactamase production of five P. shigelloides strains was characterized by activity tests, SDS-PAGE and isoelectric focusing. For all strains, MIC values of different cephalosporins were determined by microdilution methodology using inocula of 1 x 10(5) cfu/mL and 1 x 10(6) cfu/mL. Subsequently, the morphology of cells was determined by light microscopy. For one isolate, kill kinetics of cefpodoxime were determined using batch cultures with the lower and higher inocula. RESULTS: Four of five P. shigelloides strains were shown to be beta-lactamase-positive, producing different amounts of constitutively expressed non-inducible enzymes. Inoculum effects for cephalosporin susceptibility were observed for all strains. Examination of cells revealed a very strong filamentation, with filament sizes ranging from 100 microm up to 2 mm. The kill kinetics with cefpodoxime showed similar killing capacities of the antibiotic at both inoculum sizes. CONCLUSIONS: The reported resistance of P. shigelloides to cephalosporins at higher cell densities is not due to an inoculum-dependent regulation of beta-lactamases, but can be explained by the formation of extensive filaments.

Cephalosporins↗

PubMed1993

The cat/feline immunodeficiency virus model for transmucosal transmission of AIDS: nonoxynol-9 contraceptive jelly blocks transmission by an infected cell inoculum.

OBJECTIVES: To develop an animal model to study transmucosal lentivirus transmission, and to determine whether topical application of contraceptive jelly can block transmission by an infected cell incoulum. DESIGN: Feline immunodeficiency virus (FIV), a lentivirus similar to HIV, causes an AIDS-like disease in domestic cats. HIV is transmitted primarily across mucosal surfaces, and infected cells may be important in this transmission. We tested the ability of FIV-infected cells to transmit infection across the vaginal, rectal and oral mucosa of the cat, and whether a vaginal contraceptive jelly could prevent such transmission. METHODS: An inoculum consisting of 2 million FIV-infected primary cat T cells was administered vaginally, rectally or orally to female cats that had received either no pretreatment or pretreatment with a contraceptive jelly containing the detergent nonoxynol-9 as spermicide. Transmission was detected by monitoring recipient animals for viral antibodies and by viral cultures of blood leukocytes. RESULTS: A single dose of the infected cell inoculum efficiently transmitted FIV infection when delivered into the vagina or rectum (10 out of 11 animals became infected). Pretreatment of the vagina (five animals) or rectum (four animals) with contraceptive jelly protected all animals from transmission by the highly infectious inoculum. CONCLUSIONS: The cat/FIV model provides an efficient means to study transmucosal transmission of lentivirus infections, and for assessing vaginal barrier methods that could block transmission. One such method, nonoxynol-9 contraceptive jelly, effectively prevents transmucosal transmission by an FIV-infected cell inoculum.

Animals↗

PubMed1979

Effect of inoculum size on the susceptibility of Haemophilus influenzae b to beta-lactam antibiotics.

The current prevalence of ampicillin-resistant Haemophilus influenzae b meningitis requires accurate knowledge of susceptibility to alternative antibiotics. One variable affecting susceptibility is inoculum size. We studied the susceptibility of 200 clinical isolates of H. influenzae b to ampicillin, carbenicillin, and cefamandole at inocula of 10(5) and 10(7) CFU by two techniques. Fifty ampicillin-susceptible and fifty ampicillin-resistant strains were tested for susceptibility to ampicillin by broth dilution while 100 of each were tested by agar dilution. An inoculum effect was found, being greatest with the ampicillin-resistant strains. The range of minimal inhibitory concentrations for the resistant strains was 25 to 800 microgram of ampicillin per ml at an inoculum of 10(5) and 2,000 to less than 6,000 microgram of ampicillin at 10(7); 1.0 to 150 microgram of carbenicillin per ml at 10(5) and 6.2 to 2,000 microgram of carbenicillin per ml at 10(7); 0.4 to 2.0 microgram of cefamandole at 10(5) and 1.0 to 125 microgram/ml at 10(7). Because of this inoculum effect, we would not recommend the use of carbenicillin or cefamandole for therapy of ampicillin-resistant H. influenzae meningitis.

Anti-Bacterial Agents↗

PubMed1985

Reverse inoculum effect in bactericidal activity and other variables affecting killing of group B streptococci by penicillin.

Variables of the effect of penicillin G on the numbers of viable group B streptococci in broth cultures were studied. One-fourth of the MIC was the lowest concentration that reduced the viable count compared with antibiotic-free controls. The rate of killing increased with the concentration of penicillin up to 4 X MIC, but no further. During the first 2 or 3 h, the bactericidal activity was more rapid than later on. The MIC and supraoptimal concentrations of penicillin killed an inoculum of 10(6) organisms more rapidly than an inoculum of 10(4) organisms. The MIC was not inoculum dependent. The reverse inoculum effect was revealed by the killing curves but not by the MBC. There were reproducible differences among strains as to the rate of killing by penicillin; these did not correlate with the rate of multiplication, which also varied among strains. Among the 11 strains tested, there were no tolerant ones.

Female↗

PubMed1997

Antimicrobial susceptibility testing of 230 Helicobacter pylori strains: importance of medium, inoculum, and incubation time.

No standardized method of susceptibility testing for Helicobacter pylori is currently available, so before a large agar dilution study comprising 230 H. pylori strains belonging to more than 80 genetically different groups was initiated, we performed a relatively small preliminary study to determine the influences of medium, inoculum density, and incubation time. Seven media were investigated and were primarily evaluated on the basis of their abilities to support growth both semiquantitatively and qualitatively; Iso-Sensitest agar supplemented with 10% horse blood was found to be well suited for the purpose; this was closely followed by Mueller-Hinton agar with 10% horse blood, Mueller-Hinton with 10% sheep blood, and finally, 7% lysed horse blood agar. Investigations of two inoculum densities and two incubation times resulted in recommendations for the use of 10(9) CFU/ml (10[6] CFU/spot) as the inoculum and 72 h as the incubation time. A modest inoculum effect was noted for amoxicillin and metronidazole. By the methodology derived from our preliminary study, the susceptibilities of 230 H. pylori strains to six antibiotics were subsequently determined. The results were generally in accord with those of others, and apart from metronidazole, the MIC of which for approximately 25% of the strains tested was >8 microg/ml, resistance was low in Denmark. The situation might, however, quickly change when and if the number of indications for antibiotic therapy for H. pylori infections increase. Consequently, susceptibility testing of all H. pylori strains is recommended in order to survey the development of resistance, and in our hands the described methodology was relatively easy to perform and the results were easy to read.

Animals↗

PubMed2004

Impact of high-inoculum Staphylococcus aureus on the activities of nafcillin, vancomycin, linezolid, and daptomycin, alone and in combination with gentamicin, in an in vitro pharmacodynamic model.

We evaluated the impact of high (9.5 log10 CFU/g) and moderate (5.5 log10 CFU/g) inocula of methicillin-susceptible and -resistant Staphylococcus aureus (MSSA and MRSA, respectively) on the activities of nafcillin, linezolid, vancomycin, and daptomycin, alone and in combination with gentamicin in an in vitro pharmacodynamic model with simulated endocardial vegetations over 72 h. Human therapeutic dosing regimens for nafcillin, daptomycin, vancomycin, linezolid, and gentamicin were simulated. At a moderate inoculum, nafcillin (MSSA only), vancomycin, and daptomycin demonstrated equivalent and significant (P < 0.01) bactericidal (99.9% kill) activities (decreases of 3.34 +/- 1.1, 3.28 +/- 0.4, and 3.34 +/- 0.8 log10 CFU/g, respectively). Bactericidal activity was demonstrated at 4 h for nafcillin and daptomycin and at 32 h for vancomycin. Linezolid demonstrated bacteriostatic activity over the course of the study period. At a high inoculum, daptomycin exhibited bactericidal activity against both MSSA and MRSA by 24 h (decrease of 5.51 to 6.31 +/- 0.10 log10 CFU/g). Nafcillin (versus MSSA), vancomycin, and linezolid (MSSA and MRSA) did not achieve bactericidal activity throughout the 72-h experiment. The addition of gentamicin increased the rate of 99.9% kill to 8 h for daptomycin (P < 0.01) and 48 h for nafcillin (MSSA only) (P = 0.01). The addition of gentamicin did not improve the activity of vancomycin or linezolid for either isolate for the 72-h period. Overall, high-inoculum Staphylococcus aureus had a significant impact on the activities of nafcillin and vancomycin. In contrast, daptomycin was affected minimally and linezolid was not affected by inoculum.

Acetamides↗

PubMed1975

Effect of inoculum and of beta-lactamase on the anti-staphylococcal activity of thirteen penicillins and cephalosporins.

Because there are few persuasive data for selecting one semisynthetic penicillin or cephalosporin over another for treatment of serious staphylococcal infections, 118 recent clinical isolates of Staphylococcus aureus were studied to determine to what extent the presence of beta-lactamase affected the relative anti-staphylococcal activity of six penicillins and seven cephalosporins. In addition, the effect of inoculum was studied for its possible effect on the anti-staphylococcal activity of the 13 beta-lactam antibiotics. By all criteria, methicillin and nafcillin were clearly more resistant to both the inoculum effect and the production of staphylococcal beta-lactamase, whereas benzylpenicillin and cephaloridine (especially benzyl-penicillin) were the most susceptible to these effects. Cephazolin was clearly more susceptible to staphylococcal beta-lactamase and heavy inocula than the other cephalosporins (with the exception of cephaloridine), whereas cephalothin was the most resistant cephalosporin to these factors. The minimal inhibitory concentration for benzylpenicillin for tests with undiluted inoculum, compared to results with inoculum diluted 10(-4), differed by a factor up to 16,384, whereas with methicillin and nafcillin the differences were rarely more than twofold. Ratios for the other 10 antibiotics fell between these extremes. These results suggest that methicillin or nafcillin is most stable to staphylococcal beta-lactamase, and that benzylpenicillin and cephaloridine are the most susceptible.

Amidohydrolases↗

PubMed1978

Genetically marked Rhizobium identifiable as inoculum strain in nodules of soybean plants grown in fields populated with Rhizobium japonicum.

The fate of an inoculum strain of Rhizobium japonicum was studied using a genetically marked strain I-11O subline carrying resistance markers for azide, rifampin, and streptomycin (I-110 ARS). At the time of planting into a field populated with R. japonicum, seeds of soybean cultivars Kent and Peking were inoculated with varying cell densities of strain I-110 ARS. At various times during the growing season, surface-sterilized root nodules were examined for the presence of the inoculum strain by plating onto selective media. The recovery of the inoculum strain was unambiguous, varying, in the case of Kent cultivar, from about 5% with plants (sampled at 51 days) that had been inoculated with 3 X 10(8) cells per cm of row to about 20% with plants (sampled at 90 days) that had been inoculated with 3 X 10(9) cells per cm. The symbiotically incompatible interaction of Peking and strain 110 in Rhizobium-populated field soil was confirmed by the finding that at 60 days after planting, only one nodule in 360 sampled contained strain I-110 ARS. The use of genetically marked Rhizobium bacteria was found to provide for precise identification of the inoculum strain in nodules of field-grown soybeans.

Azides↗

PubMed2004

Inoculum size effect in dimorphic fungi: extracellular control of yeast-mycelium dimorphism in Ceratocystis ulmi.

We studied the inoculum size effect in Ceratocystis ulmi, the dimorphic fungus that causes Dutch elm disease. In a defined glucose-proline-salts medium, cells develop as budding yeasts when inoculated at > or = 10(6) spores per ml and as mycelia when inoculated at <10(6) spores per ml. The inoculum size effect was not influenced by inoculum spore type, age of the spores, temperature, pH, oxygen availability, trace metals, sulfur source, phosphorous source, or the concentration of glucose or proline. Similarly, it was not influenced by added adenosine, reducing agents, methyl donors, amino sugars, fatty acids, or carbon dioxide. Instead, growing cells excreted an unknown quorum-sensing factor that caused a morphological shift from mycelia to budding yeasts. This yeast-promoting effect is abolished if it is extracted with an organic solvent such as ethyl acetate. The quorum-sensing activity acquired by the organic solvent could be added back to fresh medium in a dose-dependent fashion. The quorum-sensing activity in C. ulmi spent medium was specific for C. ulmi and had no effect on the dimorphic fungus Candida albicans or the photomorphogenic fungus Penicillium isariaeforme. In addition, farnesol, the quorum-sensing molecule produced by C. albicans, did not inhibit mycelial development of C. ulmi when present at concentrations of up to 100 microM. We conclude that the inoculum size effect is a manifestation of a quorum-sensing system that is mediated by an excreted extracellular molecule, and we suggest that quorum sensing is a general phenomenon in dimorphic fungi.

Ascomycota↗

PubMed1986

Effect of inoculum size on ampicillin and amoxycillin susceptibility determined by gas-liquid chromatography for members of the family Enterobacteriaceae.

Inoculum size had only a small effect on the results for ampicillin and amoxycillin susceptibility testing of Escherichia coli. In contrast, a difference in inoculum size from 10(6) to 10(8) CFU/ml profoundly affected the results for ampicillin and amoxycillin susceptibility testing of Proteus mirabilis, causing a change from susceptibility to complete resistance. These different effects of inoculum size were observed both when susceptibility was determined by the suppression of a characteristic metabolic product analyzed by head-space gas-liquid chromatography (HS-GLC) and when it was determined by MIC testing in broth. Inoculum size affected the results for ampicillin susceptibility of P. mirabilis determined concurrently with the rapid HS-GLC urine test, because 10(8) CFU/ml may occur in urine specimens. In the rapid test, significant numbers of Escherichia, Klebsiella, Citrobacter, Proteus, Morganella, and Providencia spp. in urine specimens are detected within 4 h by HS-GLC analysis for characteristic metabolic products in cultures. Most P. mirabilis in urine specimens appeared to be ampicillin resistant in the rapid HS-GLC test but were reported to be ampicillin susceptible in hospital laboratory agar dilution tests 2 days later. However, ampicillin susceptibility results for Escherichia, Klebsiella, and Citrobacter spp. agreed with hospital laboratory reports. It was concluded that reports of ampicillin susceptibility or resistance for the commonest cause of urinary tract infections, E. coli, within 4 h of receipt of the specimen would be clinically valuable and that a provisional report of ampicillin resistance for P. mirabilis would not lead to ineffective therapy.

Amoxicillin↗

PubMed1985

Influence of inoculum size, incubation temperature, and cell culture density on virus detection in environmental samples.

The influence of inoculum size and cell culture density on virus titer by cytopathic effect or plaque assay was studied using poliovirus type 1 and BGM (Buffalo green monkey) cells as a model for this evaluation. With a plaque assay system, a linear relationship was observed for an inoculum size of up 1 mL/25 cm2; a marked decrease in the number of plaques was observed when over 1 mL of sample was inoculated on this surface area. Cell culture density also affected virus titer; maximal titers were observed when cells were seeded at 25 000 to 75 000 cells/mL and incubated for 6 days before infection with the virus. Viral density, evaluated as most-probable-number and measured by cytopathic effect under liquid overlay, revealed that the viral titer was similar up to 1 mL inoculum and increased only when over 1 mL was inoculated. Cell density had no significant effect on the viral titer measured by the most-probable-number method and cytopathic effect. Inactivation of inoculum due to an incubation temperature of 37 degrees C for a short period was shown to be minimal for poliovirus type 1, reovirus type 2, coxsackievirus B-5, and the simian rotavirus SA-11. Longer inactivation time led to a 2 logs reduction of the infectious titer of coxsackievirus B-5 (in 48 h) while the other viruses showed a significant reduction in titer only after 96 h.

Animals↗

PubMedDate not supplied

Degradation of oleic acid in anaerobic filters: the effect of inoculum acclimatization and biomass recirculation.

The degradation of oleic acid in anaerobic filters was studied and the effect of an acclimated inoculum and biomass recirculation was evaluated. Three anaerobic filters (R1, R2, and R3) were operated in parallel. The anaerobic filters R1 and R2 were inoculated with nonacclimated biomass, whereas the anaerobic filter R3 was inoculated with acclimated biomass. In the anaerobic filters R2 and R3, biomass settling and recirculation were applied. The use of an acclimated inoculum and biomass recirculation (R3) was beneficial in terms of removal efficiency, which was 4 to 8% higher than in the anaerobic filters R1 and R2 when oleate was the sole carbon source fed to the reactors at an applied organic load of 12.5 kg of chemical oxygen demand (COD)/m3 x d, even with an oleate to calcium and magnesium ion molar concentration ratio of 6.8. Biomass recirculation significantly reduced the biomass washout and the toxic effect on the acetogenic and methanogenic populations. The use of an acclimated inoculum was beneficial in terms of methane yield, which was 50% greater than that observed for the reactors inoculated with nonacclimated inoculum for the highest applied organic loading rate (12.5 kg COD/m3 x d). At the end of the operation, the biomass was encapsulated by a whitish matter, which was well detected by microscopic examination. When this sludge was incubated in batch vials at 37 degrees C where no substrate was added, methane production from the adsorbed organic matter was evidenced, attaining a maximum value (at standard temperature and pressure) of 39.7 mL/g volatile solids x d for the biomass taken from R1. With stirring (150 r/min), the methane production rate was 13.8 times higher than under static conditions. When oleate was added to this sludge, methane production was delayed, suggesting that adsorbed matter can be an intermediate of oleate degradation such as stearic, palmitic, myristic, or other saturated acids.

Anaerobiosis↗

PubMed2003

Effect of inoculum size, relative humidity, storage temperature, and ripening stage on the attachment of Salmonella Montevideo to tomatoes and tomatillos.

The influence of inoculum populations and environmental factors on attachment of Salmonella Montevideo to the surface of tomatoes and tomatillos was evaluated. To study the effect of inoculum size, red, ripe tomatoes were spot-inoculated with bacterial suspensions (10(5) and 10(8) CFU/fruit) and stored at 22 degrees C under 100% relative humidity. The effects of temperature (12, 22, and 30 degrees C) and relative humidity (75, 85, and 97%) on attachment of the pathogen (10(7) CFU/fruit) to tomatoes (red and green) and ripe tomatillos were also evaluated. Inoculated fruits were stored for 90 min at all combinations of temperature and relative humidity, and after rinsing with water, the number of cells attached to the surface was determined. Salmonella Montevideo attached to the surface of tomatoes within 90 min. A direct correlation between the number of attached cells and the population in the inoculum was observed. The percentage of cells that attached immediately after inoculation was approximately 0.3% for the three test products. After storage for 90 min at various temperature and relative humidity conditions, the number of adhering cells ranged from 4.0 to 5.4 log CFU/fruit (1.2% of inoculum). Both the type of product and the temperature/relative humidity combination had a significant (P < 0.05) effect on attachment of Salmonella Montevideo to the surfaces of tomatoes and tomatillos. Scanning electron micrographs of the cuticles of inoculated washed tomatoes and tomatillos revealed typical skin cell patterns, and only a few randomly dispersed Salmonella Montevideo were observed. Deposition of Salmonella Montevideo on the surface of tomatoes and tomatillos could result in attachment and subsequent colonization under suitable conditions.

Bacterial Adhesion↗

PubMed2004

Attachment of Salmonella Poona to cantaloupe rind and stem scar tissues as affected by temperature of fruit and inoculum.

A negative temperature differential between fruits or vegetables and the water in which they are immersed theoretically enhances infiltration of water and any microorganisms it might contain into tissues. The effect of temperature differentials between cantaloupes and wash water, each at 4 and 30 degrees C, on changes in cantaloupe weight and populations of Salmonella enterica Poona recovered from rinds and stem scar tissues of Eastern and Western (shipper) types of cantaloupes was assessed. The percent weight increase in Western cantaloupes was significantly greater (P < or = 0.05) than that in Eastern cantaloupes for all cantaloupe and inoculum temperature combinations. Salmonella Poona attachment to or infiltration of Eastern but not Western cantaloupe rind is enhanced when the fruit is at 4 degrees C, compared with 30 degrees C, regardless of the temperature of the immersion suspension. The number of Salmonella Poona cells recovered from rind tissue of Western cantaloupes at 30 degrees C immersed in inoculum at 30 degrees C was significantly less (P < or = 0.05) than that recovered from rind tissues of cantaloupes at 4 or 30 degrees C that were immersed in inoculum at 4 degrees C. Salmonella Poona in immersion water can adhere to or infiltrate surface tissues of cantaloupes. The populations of Salmonella Poona recovered from stem scar tissues of Eastern and Western types of cantaloupes were not significantly (P > 0.05) affected by cantaloupe and inoculum temperature combinations. Populations of cells adhering to or infiltrating various cantaloupe tissues is not dictated entirely by temperature differentials between fruits and immersion suspensions: rather, it also apparently is influenced by structures unique to surface tissues.

Bacterial Adhesion↗