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At least 415 records · Page 23Linked to original sources
PubMed1999

In vitro maturation of human preovulatory oocytes reconstructed by germinal vesicle transfer.

OBJECTIVE: To describe a micromanipulation-electrofusion procedure for transferring germinal vesicles (GVs) between immature human oocytes. DESIGN: Pilot study to assess oocyte maturation after an invasive micromanipulation procedure. SETTING: Research laboratory at a university medical center. PATIENT(S): Immature oocytes were discarded from intracytoplasmic sperm injection (ICSI)-IVF cycles of patients 23-48 years of age. INTERVENTION(S): Initially, GV removal and transfer were performed on the same oocyte; these "self-reconstructed" oocytes were then cultured in vitro for up to 50 hours and examined periodically for maturation as judged by the extrusion of the first polar body. In a second study, GVs from oocytes of "old" patients (>38 years old) were successfully transferred into enucleated immature oocytes of "young" patients (<31 years old). MAIN OUTCOME MEASURE(S): Extrusion of the first polar body was monitored in "reconstructed" and control oocytes; karyotypes also were analyzed at meiosis II. RESULT(S): From 48 oocytes from old patients, 12 GVs were successfully removed, transferred, and fused into previously enucleated oocytes from young patients. After in vitro culture, 7 of these "reconstructed" oocytes matured to meiosis II, a maturation rate not significantly different from that observed in nonmanipulated controls. A normal, second meiotic metaphase chromosome complement was observed in 4 of 5 reconstructed oocytes. CONCLUSION(S): Normal meiosis can occur after the transfer of a GV into an enucleated host oocyte. Germinal vesicle transfer may be a valuable research procedure that generates cell models to characterize the cytoplasmic-nuclear interplay for cell cycle regulation, maturation, and fertilization in the human oocyte; it also may be a potentially attractive alternative to oocyte donation.

Adult↗
PubMed2003

Micromechanical studies of mitotic chromosomes.

We review micromechanical experiments studying mechanoelastic properties of mitotic chromosomes. We discuss the history of this field, starting from the classic in vivo experiments of Nicklas (1983). We then focus on experiments where chromosomes were extracted from prometaphase cells and then studied by micromanipulation and microfluidic biochemical techniques. These experiments reveal that chromosomes have a well-behaved elastic response over a fivefold range of stretching, with an elastic modulus similar to that of a loosely tethered polymer network. Perturbation by microfluidic "spraying" of various ions reveals that the mitotic chromosome can be rapidly and reversibly decondensed or overcondensed, i.e., that the native state is not maximally compacted. We compare our results for chromosomes from cells to results of experiments by Houchmandzadeh and Dimitrov (1999) on chromatids reconstituted using Xenopus egg extracts. Remarkably, while the stretching elastic response of reconstituted chromosomes is similar to that observed for chromosomes from cells, reconstituted chromosomes are far more easily bent. This result suggests that reconstituted chromatids have a large-scale structure that is quite different from chromosomes in somatic cells. Finally, we discuss microspraying experiments of DNA-cutting enzymes, which reveal that the element that gives mitotic chromosomes their mechanical integrity is DNA itself. These experiments indicate that chromatin-condensing proteins are not organized into a mechanically contiguous "scaffold," but instead that the mitotic chromosome is best thought of as a cross-linked network of chromatin. Preliminary results from restriction enzyme digestion experiments indicate a spacing between chromatin "cross-links" of roughly 15 kb, a size similar to that inferred from classical chromatin loop isolation studies. These results suggest a general strategy for the use of micromanipulation methods for the study of chromosome structure.

Animals↗
PubMed1999

Micrometer dimension derivatization of biosensor surfaces using confocal dynamic patterning.

Using laser scanning confocal optics in conjunction with avidin/biotin technology, micrometer-sized patterns of biomolecules were fabricated on glassy-carbon and fused-silica surfaces. Photoactive biotin was immobilized using the 325-nm line of a Helium-Cadmium laser, which was focused through a 25x or 100x quartz microscope objective. A three-dimensional piezoelectric micromanipulator was used to position the sample surface in the focal plane of the microscope objective and to create patterns on the focused surface. Biotin patterns with line widths of 5-20 microns were produced by varying the scan speed of the micromanipulator while exposing the surface to the laser. The integrity of the immobilized biotin was confirmed by subsequent derivatization with fluorescently labeled avidin. Fluorescence microscopy with a cooled charge coupled device (CCD) imaging system was used to visualize the distribution of biotin and fluorescent avidin within the patterns created by the laser.

Biosensing Techniques↗
PubMed2000

Pore sizes in hydrated dextran microspheres.

The average pore size of hydrated dextran microspheres is derived from rheological and protein release data. The microspheres were prepared by cross-linking an aqueous solution of methacrylated dextran emulsified in a continuous poly(ethylene glycol) phase. The rheological data were obtained using a novel micromanipulation technique, which enables the compression of a single microsphere. The so obtained pseudoelasticity moduli of the microspheres were derived from these compression data and corresponded well with the elasticity moduli of macroscopic hydrogels of the same composition, as determined with dynamic mechanical analysis. The modulus increased with decreasing water contents of the microspheres and with increasing degrees of methacrylate substitution of the dextran used. Furthermore, the average pore sizes calculated from the pseudoelasticity moduli were in good agreement with the pore sizes derived from protein release data. In conclusion, this study shows that micromanipulation provides insight into the average pore sizes of dextran microspheres, which is an important characteristic that will modulate the release of encapsulated proteins.

Dextrans↗
PubMed2003

Manipulated mouse embryos as bioassay system for water quality control.

Mouse pronuclear stage embryos with intact slit zona pellucida (manipulated) were cultured in vitro until the hatched blastocyst stage in simplex optimized medium with higher K+ concentration (KSOM) prepared with three different water types: tap, deionized reverse osmosis (D-O) water and Milli-Q system (M-Q) water. The culture media were supplemented with or without protein and ethylenediaminetetraacetic acid (EDTA, disodium salt). The rates of hatched blastocysts were significantly affected (p < 0.01) by micromanipulation, protein supplement and water source. The water source has no influence (p > 0.05) on development in EDTA-supplemented protein-free culture media, whereas in EDTA- and protein-free culture media, the water quality significantly (p < 0.001) affected the rates of development, with higher rates in media prepared with M-Q water. The micromanipulated embryos showed higher sensitivity to the water quality (p < 0.01). It worth mentioning that the rates of hatched blastocysts in protein-free culture media were very low (0-7.5%). Furthermore, the three different water types were analysed by measuring the electrical conductivity, inorganic ions, total organic carbon and endotoxins to evaluate the purity. M-Q water showed the lowest levels of inorganic ion, total organic carbon and endotoxin concentrations. We concluded that manipulated mouse embryos are good system to evaluate the quality of water used in biological system.

Animals↗
PubMed2003

A comparison of established and new approaches in ovine and bovine nuclear transfer.

Several breakthroughs in nuclear transfer research were first achieved in sheep, although cattle soon became the main livestock species of interest. However, sheep still offer significant advantages both in basic and applied research. With increased interest in cloning of livestock, new approaches have been developed for both sheep and cattle nuclear transfer technology. These include methods for zona-free nuclear transfer that can be performed with or without the use of micromanipulator. Here we describe four different nuclear transfer methods including the traditional micromanipulation-assisted method in sheep, zona-free method in sheep in which the order of enucleation and nucleus delivery have been reversed ("reverse-order" cloning) and zona free manual cloning methods ("hand-made cloning") for embryonic and somatic cloning in cattle. The purpose of this paper is to encourage people to familiarize themselves with these different methods available and to help them choose and test the method most suitable for their particular circumstances.

Animals↗
PubMed1996

Birth of normal mice after removal of the supernumerary male pronucleus from polyspermic zygotes.

Each year, world wide, tens of thousands of zygotes derived from the in-vitro insemination of human oocytes undergo polyspermic fertilization. These embryos must presently be discarded because it has never been demonstrated in any mammal that polyspermic zygotes can develop normally to term after removal of the supernumerary male pronucleus. Our study was undertaken to test the developmental potential of polyspermic zygotes corrected by micromanipulation. Mouse oocytes were inseminated zona-free, and polyspermic zygotes were manipulated so as to remove one of the two male pronuclei. Surviving embryos were then observed for further development in vitro and after transfer into pseudopregnant females. Of 58 zygotes manipulated, 18 developed to the blastocyst stage and were transferred. Five animals (two male and three females) were born. The agouti coat colour marker confirmed the genotypes of the gametes. All five animals developed to normal-appearing adults, and all five produced at least 10 normal offspring. One adult founder animal was karyotyped and found to have a normal chromosome complement. These results demonstrate for the first time that a mammalian egg that has undergone polyspermic fertilization can develop normally after restoration of the diploid state by micromanipulation. Accordingly, the results provide impetus for attempting to rescue polyspermic human embryos.

Animals↗
PubMed1997

Assisted hatching by zona drilling of human embryos in women of advanced age.

The study was conducted to examine the efficacy of the zona drilling technique for promotion of successful implantation and pregnancy by assisting embryo hatching in women of advanced age undergoing treatment by in-vitro fertilization. A total of 839 embryos from 211 patients aged >38 years underwent assisted hatching during 312 cycles of therapy. The outcome of this micromanipulation procedure was compared to 540 non-hatched pre-embryos transferred to 174 patients during 274 cycles of therapy. Assisted hatching was performed on four- to eight-cell stage embryos using the zona drilling technique. In the assisted hatching group, 839 micromanipulated embryos were replaced (two to four embryos per patient) compared to 540 embryos transferred in the control group. Despite the fact that the pregnancy rate was not statistically different between the groups (8.9% in the assisted hatching group versus 5.1% in the controls) a trend towards an increase was noted in the assisted hatching group. The implantation rate was 3.75 and 3.55% per patient respectively, and there was no significant difference in abortion rate between the groups. The delivery rate was 3.8 and 3.4% per cycle respectively. The results of this study demonstrated that assisted hatching by zona drilling in a selected group of patients aged >38 years does not increase the take-home baby rate after in-vitro fertilization treatment.

Adult↗
PubMed1992

Implantation enhancement by selective assisted hatching using zona drilling of human embryos with poor prognosis.

Assisted hatching by zona drilling using acidic Tyrode's solution was performed during three randomized trials in 330 in-vitro fertilization patients. The trials were designed in order to study the overall effect of the procedure and whether characteristic patient [i.e. maternal age and basal levels of follicle stimulating hormone (FSH)] and embryonic features (i.e. zona pellucida thickness) are important for the decision to perform assisted hatching routinely. Couples (n = 137; Trial 1) in whom the female partner had normal basal FSH levels were randomized in a control group (without micromanipulation) and a zona drilling group (all embryos micromanipulated). The incidence of implantation (67/239; 28%) of zona-drilled embryos compared favourably with that of control embryos (49/229; 21%), but the difference was not significant. Retrospective analysis revealed that those embryos whose zonae were thicker than 15 microns were rescued. In order to test the validity of this finding, selective assisted hatching was performed on embryos with a poor prognosis in 163 other patients (Trial 2). The couples were randomized into a control group and a group in which embryos were selectively zona-drilled, based on zona thickness and other embryonic features. The rate of embryonic implantation in the selectively zona-drilled group was 25% (70/278), significantly (P less than 0.05) higher than that of control embryos (51/285; 18%). Although it was demonstrated retrospectively and prospectively that assisted hatching by zona drilling is effective in embryos with thick zonae (greater than or equal to 15 microns), patients whose embryos have thin (less than 13 microns) zonae may be jeopardized by the procedure.(ABSTRACT TRUNCATED AT 250 WORDS)

Embryo Implantation↗
PubMed2003

Handmade somatic cell cloning in cattle: analysis of factors contributing to high efficiency in vitro.

Widespread application of somatic cell cloning has been hampered by biological and technical problems, which include complicated and time-consuming procedures requiring skilled labor. Recently, zona-free techniques have been published with limited or no requirement for micromanipulators. The purpose of the present work was to optimize certain steps of the micromanipulator-free (i.e., handmade) procedure, to analyze the morphology of the developing blastocysts, and to explain factors involved in the high efficiencies observed. Optimization of the procedure included selection of the appropriate medium for enucleation, orientation of pairs at fusion, timing of fusion, and culture conditions. As a result of these improved steps, in vitro efficiency as measured by blastocysts per reconstructed embryo and blastocysts per working hour was among the highest described so far. The cattle serum used in our experiments was superior to other protein sources for in vitro embryo development. One possible explanation of this effect is the considerable mitogenic activity of the cattle serum compared with that of commercially available fetal calf serum. Morphological analysis of blastocysts by inverted microscopy, inner cell mass-trophoblast differential staining, and transmission electron microscopy revealed high average quality. A high initial pregnancy rate was achieved after the transfer of single blastocysts derived by aggregation of two nuclear transfer embryos into recipients. The improved handmade somatic cell nuclear transfer method may become a useful technology as a simple, inexpensive, and efficient alternative to traditional somatic cell nuclear transfer.

Animals↗
PubMed1995

Enucleation by centrifugation of in vitro-matured bovine oocytes for use in nuclear transfer.

Nuclear transfer has the potential to produce large numbers of identical progeny. Current limitations of the technique are associated with the use of micromanipulation for the demanding enucleation and reconstitution procedure. With the overcoming of this limitation, increased numbers of nuclear transfer embryos could be produced. Centrifugation of bovine oocytes at 15,000 x g for 2 min resulted in the stratification of organelles within the cytoplasm which positioned the metaphase II spindle for enucleation. After removal of the zona pellucida with Pronase, the oocytes were centrifuged in a Percoll density gradient so that the oocytes were stretched apart to form cytoplasts and the metaphase II spindle was separated from the majority of oocytes. Enucleation by centrifugation efficiently produced a consistent population of enucleated cytoplasts from bovine in vitro-matured oocytes. The population of enucleated cytoplasts was enriched by exclusion of the cytoplasts that exhibited an extrusion cone containing metaphase II chromosomes 6 h after centrifugation. The enucleated oocyte cytoplasts were aggregated with blastomeres isolated from in vivo-collected morulae. The aggregated embryonic cells were electrofused to obtain nuclear transfer embryos that were placed into a sodium alginate false zona and were capable of cleavage and development in vitro. The development of nuclear transfer embryos produced through use of centrifugation and aggregation techniques was comparable with that of nuclear transfer embryos produced by micromanipulation techniques.

Alginates↗
PubMed1967

Large bodies of Mycoplasma and L-form organisms.

The large bodies of various Mycoplasma and L-form organisms were studied by ultraviolet fluorescence microscopy of preparations stained with various fluorochromes. Primuline and Thioflavine S specifically stained the outer portion or rim of the large bodies, and the fluorescence characteristics of the stained bodies differed from those for other microorganisms and for spheroplasts and protoplasts. Small granular structures similar in size and morphology to minimal reproductive units were observed within some of the large bodies by phase microscopy and by fluorescence microscopy with acridine orange or Coriphosphine O. Micromanipulation probing of the large bodies revealed their elastic nature; many of the large bodies could be subdivided into two or more smaller circular bodies, each retaining the fluorescence staining properties of the parent body. Under these conditions, however, a few of the large bodies were ruptured, leaving the stainable outer boundary area as a stable residual structure. The large bodies were somewhat resistant to various rigorous treatments normally employed to eliminate viability of Mycoplasma and L-form cultures. Structures similr to large bodies were observed in various natural tissues, and structures resembling large bodies in size, morphology, fluorescence staining characteristics, and reaction to micromanipulation probing were reconstructed from an acetone extract of egg yolk. Overall, the large bodies of Mycoplasma and L-form organisms appeared to be structures resulting from accumulations of metabolic by-products and medium components within or on which minimal reproductive units had become entrapped, although it could not be ruled out that they might be defined structures specifically formed during culture as protective lipoidal sacs for the minimal reproductive units.

Bacteria↗
PubMed1997

Clinical review 87: In vitro fertilization for male factor infertility.

Since the first U.S. report of a successful delivery from in vitro fertilization in 1982 (65), progress in the field of assisted reproduction and micromanipulation has been truly dramatic. Perhaps the most exciting advances have been in the area of male factor infertility. Couples who previously would have been offered donor insemination or adoption are now achieving pregnancies despite severe impairments in semen quality, the presence of only single numbers of sperm in the ejaculate, or unreconstructable reproductive tract obstruction. Techniques of micromanipulation that were revolutionary less than five yr ago are now obsolete, replaced by even more successful methods. Even nonobstructive azoospermia resulting from maturation arrest or other impairments in germ cell development have been added to the list of treatable factors in male infertility, as sperm can frequently be extracted directly from testicular parenchyma that is aspirated or surgically biopsied. For patients without sperm in the testicular parenchyma, round spermatid or secondary spermatocyte injections are at least theoretically possible. Several important questions remain with regard to IVF-ICSI. 1) What should be the specific indications for IVF and IVF-ICSI? Should IVF alone ever be used for male factor infertility? 2) What are the reasons for failure to achieve pregnancy after ICSI, which still represent over half of our attempts at achieving ongoing pregnancies? 3) Can we be certain that using severely impaired or less mature sperm will not result in significant birth defects or in genetic abnormalities that could affect the offspring in adolescence or adulthood? 4) What is the most successful and cost effective approach for the infertile couple with impaired semen parameters? For couples with male factor infertility, careful evaluation and treatment of the man should be considered before assisted reproduction, including ICSI. Contemporary application of ICSI for severe male factor infertility can allow pregnancy rates up to 52% (33), with ongoing pregnancy and live delivery rates as high as 37% per IVF cycle attempt (27). As long as viable sperm are present in the ejaculate or retrievable from the reproductive tract, then ICSI procedures can be applied.

Congenital Abnormalities↗
PubMed1984

Production of monozygotic mouse twins from microsurgically bisected morulae.

Mouse monozygotic twins were produced by bisection of the compacted morulae and transfer of the pairs of half-embryos after culture in vitro. The compacted morulae (about 16 cells) were microsurgically bisected, using a fine glass needle attached to a micromanipulator, without any supporting micro-instruments, after pretreatment for zona-softening and decompaction. About 80% of the morulae were bisected without visible cell damage. After 20 h in culture, the half-embryos were classified morphologically as eu-blastocysts, pseudo-blastocysts, or trophectodermal vesicles or non-integrated forms. After culture of 131 pairs of bisected morulae, 75 (57.3%) pairs of eu-blastocysts, 20 (15.3%) pairs comprising a eu-blastocyst and pseudo-blastocyst, and 9 (6.9%) pairs of pseudo-blastocysts, were obtained. The pseudo-blastocysts were considered to be derived from half-morulae in which some blastomeres were destroyed or dissociated as a result of micromanipulation. From 30 pairs of eu-blastocysts transferred to 21 recipients, 5 twin fetuses on Day 17 (18 pairs/9 recipients) and 3 twin male young (12 pairs/12 recipients) were obtained. Survival rate of the twin-embryo pairs was 27.8% at autopsy and 25.0% at term. None of the 20 pairs of pseudo-blastocysts transferred to 10 recipients gave rise to normal conceptuses.

Animals↗
PubMed2001

Refrigeration of donor cells in preparation for bovine somatic nuclear transfer.

In mammals, preparation of donor cells for somatic nuclear transfer is very important because the character of the donor cell directly affects the efficiency and outcome of transfer. The protocols used most commonly for donor preparation are (i) disaggregating cells from fresh tissue 1-2 h before micromanipulation or (ii) trypsinizing cultured cells temporarily, after special treatments for 3-8 days (for example, serum starvation). In this study, a new simple protocol was designed, whereby the donor cells (cumulus cells) used in bovine somatic nuclear transfer were refrigerated. In brief, cultured cells at 80-100% confluency were detached using trypsin, washed by centrifugation, aliquoted into different vials and refrigerated at 4 degrees C. The density of viable cells was decreased after day 1 of refrigeration; however, the rate of decrease tended to slow down with increasing duration of refrigeration. Cells refrigerated for 15 days were seeded at a density of 5 x 10(4) ml(-1) and reached 70% confluency after day 2 of culture. Most cells had the normal number of chromosomes (2n = 60). Cells chilled at 4 degrees C for different durations were removed from refrigeration and immediately subjected to micromanipulation. The in vitro development of reconstructed embryos (fusion rates, cleavage rates, morula and blastocyst rates) indicated that there were no significant differences among treatment groups regardless of the duration of refrigeration (0-2 weeks) of the donor cells. Reconstructed embryos were transferred into the uteri of recipient cows. No significant differences were observed in established early pregnancies between embryos derived from the non-refrigerated donor cells and those derived from refrigerated donor cells. This study indicates that refrigeration of donor cells for 1-2 weeks is a feasible protocol for preparing donor cells for bovine somatic nuclear transfer, and does not compromise development in vitro and early development in vivo.

Animals↗
PubMed2002

Micromechanical studies of mitotic chromosomes.

We review micromechanical experiments on mitotic chromosomes. We focus on work where chromosomes were extracted from prometaphase amphibian cells, and then studied by micromanipulation and microfluidic biochemical techniques. These experiments reveal that chromosomes have well-behaved elastic response over a fivefold range of stretching, with an elastic modulus similar to that of a loosely tethered polymer network. Perturbation by microfluidic 'spraying' of various ions reveals that the mitotic chromosome can be rapidly and reversibly decondensed or overcondensed, i.e. that the native state is not maximally compacted. Finally, we discuss microspraying experiments of DNA-cutting enzymes which reveal that the element which gives mitotic chromosomes their mechanical integrity is DNA itself. These experiments indicate that chromatin-condensing proteins are not organized into a mechanically contiguous 'scaffold', but instead that the mitotic chromosome is best thought of as a cross-linked network of chromatin. Preliminary results from restriction-enzyme digestion experiments indicate a spacing between chromatin 'cross-links' of roughly 15 kb, a size similar to that inferred from classical chromatin-loop-isolation studies. We compare our results to similar experiments done by Houchmandzadeh and Dimitrov (J Cell Biol 145: 215-213 (1999)) on chromatids reconstituted using Xenopus egg extracts. Remarkably, while the stretching elastic response of the reconstituted chromosomes is similar to that observed for chromosomes from cells, the reconstituted chromosomes are far more easily bent. This result suggests that reconstituted chromatids have a large-scale structure which is quite different from chromosomes in somatic cells. More generally our results suggest a strategy for the use of micromanipulation methods for the study of chromosome structure.

Animals↗
PubMed1994

Preimplantation development of manipulated mouse zygotes fused with the second polar bodies: a cytogenetic study.

Immediately after fertilization one chromatid of each maternally-derived chromosome is extruded into the second polar body (2PB). We tested the ability of these "extra" chromosomes to support preimplantation development. Micromanipulation and electrofusion techniques were used to fuse 2PBs with diploid, haploid, or enucleated mouse zygotes. Androgenetic haploids, intact embryos, and digynic triploids served as the controls for the reconstructed embryos. Androgenetic haploid zygotes developed to the blastocyst stage only when fused with the 2PBs. This result demonstrates that even when extruded into the 2PB, chromosomes retain their ability to support normal preimplantation development. However, 2PB fusion with diploid zygote impaired preimplantation development. Normal development of experimentally produced digynic triploids (zygotes with one extra maternal pronucleus) indicated that developmental arrests, caused by the 2PB fusion, were not the result of triploidy or micromanipulation procedures. Cytogenetic studies showed that developmental failures of the reconstructed embryos were caused by premature chromosome condensation of the polar body chromosomes. This result indicates that 2PB must be removed from the zygotes' perivitelline space during animal cloning experiments. In addition, we showed that 2PB fusion with enucleated zygote is a reliable method for 2PB karyotyping and may be used in the studies of mammalian meiosis.

Aneuploidy↗
PubMed1998

Laser tweezers and optical microsurgery in cellular and molecular biology. Working principles and selected applications.

After focusing in a microscope, light can be used for micromanipulation of (sub-)micrometer sized objects. Focused beams of classical light ablate elements of the cell division machinery and switch the beating of hearts on a cellular basis. Focused lasers (laser microbeams or optical scissors) allow in addition very precise nanomachining in a wide field of applications, from developmental biology to plant biotechnology. While in microbeam work high power densities and efficient light-tissue interactions are required, optical tweezers work in a complementary way: Moderate power densities and small absorption of the laser by the biological material is needed. With light pressure and optical gradient forces optical tweezers can be used to move microscopic objects, even in the interior of closed cells. In total, most mechanical micromanipulation techniques known from cellular and molecular biology can be replaced by their optical correlate and some applications are possible which cannot be performed micromechanically. When these optical microtools are operated at their maximum performance, the physical effects are as interesting as their biological applications: The laser microbeam can generate extreme local temperatures, which however are dissipated within a few tens of nanoseconds and therefore cause damage only very locally. The optical tweezers with a working wavelength in the optical window of biological material (600-1100 nm) exert piconewton forces without any mechanical contact. The present article discusses some quantitative physical aspects of microbeams and optical tweezers and gives a few selected examples of applications.

Animals↗