Nonlinear Ringdown at the Black Hole Horizon.

The gravitational waves emitted by a perturbed black hole ringing down are well described by damped sinusoids, whose frequencies are those of quasinormal modes. Typically, first-order black hole perturbation theory is used to calculate these frequencies. Recently, it was shown that second-order effects are necessary in binary black hole merger simulations to model the gravitational-wave signal observed by a distant observer. Here, we show that the horizon of a newly formed black hole after the head-on collision of two black holes also shows evidence of nonlinear modes. Specifically, we identify one quadratic mode for the l=2 shear data, and two quadratic ones for the l=4, 6 data in simulations with varying mass ratio and boost parameter. The quadratic mode amplitudes display a quadratic relationship with the amplitudes of the linear modes that generate them.

Multimode Quasinormal Spectrum from a Perturbed Black Hole.

When two black holes merge, the late stage of gravitational wave emission is a superposition of exponentially damped sinusoids. According to the black hole no-hair theorem, this ringdown spectrum depends only on the mass and angular momentum of the final black hole. An observation of more than one ringdown mode can test this fundamental prediction of general relativity. Here, we provide strong observational evidence for a multimode black hole ringdown spectrum using the gravitational wave event GW190521, with a maximum Bayes factor of 56±1 (1σ uncertainty) preferring two fundamental modes over one. The dominant mode is the ℓ=m=2 harmonic, and the subdominant mode corresponds to the ℓ=m=3 harmonic. The amplitude of this mode relative to the dominant harmonic is estimated to be A_{330}/A_{220}=0.2_{-0.1}^{+0.2}. We estimate the redshifted mass and dimensionless spin of the final black hole as 330_{-40}^{+30}M_{⊙} and 0.86_{-0.11}^{+0.06}, respectively. We find that the final black hole is consistent with the no-hair theorem and constrain the fractional deviation from general relativity of the subdominant mode's frequency to be -0.01_{-0.09}^{+0.08}.

News from Horizons in Binary Black Hole Mergers.

In a binary black hole merger, it is known that the inspiral portion of the waveform corresponds to two distinct horizons orbiting each other and that the merger and ringdown signals correspond to the final horizon being formed and settling down to equilibrium. However, we still lack a detailed understanding of the relation between the horizon geometry in these three regimes and the observed waveform. Here we show that the well-known inspiral chirp waveform has a clear counterpart on black hole horizons, namely, the shear of the outgoing null rays at the horizon. We demonstrate that the shear behaves very much like a compact binary coalescence waveform with increasing frequency and amplitude. Furthermore, the parameters of the system estimated from the horizon agree with those estimated from the waveform. This implies that even though black hole horizons are causally disconnected from us, assuming general relativity to be true, we can potentially infer some of their detailed properties from gravitational wave observations.

Interior of a Binary Black Hole Merger.

We find strong numerical evidence for a new phenomenon in a binary black hole spacetime, namely, the merger of marginally outer trapped surfaces (MOTSs). By simulating the head-on collision of two nonspinning unequal mass black holes, we observe that the MOTS associated with the final black hole merges with the two initially disjoint surfaces corresponding to the two initial black holes. This yields a connected sequence of MOTSs interpolating between the initial and final state all the way through the nonlinear binary black hole merger process. In addition, we show the existence of a MOTS with self-intersections formed immediately after the merger. This scenario now allows us to track physical quantities (such as mass, angular momentum, higher multipoles, and fluxes) across the merger, which can be potentially compared with the gravitational wave signal in the wave zone, and with observations by gravitational wave detectors. This also suggests a possibility of proving the Penrose inequality mathematically for generic astrophysical binary back hole configurations.

Seeing is believing: the impact of RB on nuclear organization.

The retinoblastoma tumor suppressor (RB) prevents G1 to S cell cycle transition by inhibiting E2F activity. This function requires that RB remains un- or underphosphorylated (the so-called active forms of RB). Recently, we showed that active forms of RB cause widespread changes in nuclear architecture that are visible under a microscope. These phenotypes did not correlate with cell cycle arrest or repression of the E2F transcriptional program, but appeared later, and were associated with the appearance of autophagy or in IMR-90 cells with senescence markers. In this perspective, we describe the relative timing of these RB-induced events and discuss the mechanisms that may underlie RB-induced chromatin dispersion. We consider the relationship between RB-induced dispersion, autophagy, and senescence and the potential connection between dispersion and cell cycle exit.

Active RB causes visible changes in nuclear organization.

RB restricts G1/S progression by inhibiting E2F. Here, we show that sustained expression of active RB, and prolonged G1 arrest, causes visible changes in chromosome architecture that are not directly associated with E2F inhibition. Using FISH probes against two euchromatin RB-associated regions, two heterochromatin domains that lack RB-bound loci, and two whole-chromosome probes, we found that constitutively active RB (ΔCDK-RB) promoted a more diffuse, dispersed, and scattered chromatin organization. These changes were RB dependent, were driven by specific isoforms of monophosphorylated RB, and required known RB-associated activities. ΔCDK-RB altered physical interactions between RB-bound genomic loci, but the RB-induced changes in chromosome architecture were unaffected by dominant-negative DP1. The RB-induced changes appeared to be widespread and influenced chromosome localization within nuclei. Gene expression profiles revealed that the dispersion phenotype was associated with an increased autophagy response. We infer that, after cell cycle arrest, RB acts through noncanonical mechanisms to significantly change nuclear organization, and this reorganization correlates with transitions in cellular state.

Dynamic and Stable Cohesins Regulate Synaptonemal Complex Assembly and Chromosome Segregation.

Assembly of the synaptonemal complex (SC) in Drosophila depends on two independent pathways defined by the chromosome axis proteins C(2)M and ORD. Because C(2)M encodes a Kleisin-like protein and ORD is required for sister-chromatid cohesion, we tested the hypothesis that these two SC assembly pathways depend on two cohesin complexes. Through single- and double-mutant analysis to study the mitotic cohesion proteins Stromalin (SA) and Nipped-B (SCC2) in meiosis, we provide evidence that there are at least two meiosis-specific cohesin complexes. One complex depends on C(2)M, SA, and Nipped-B. Despite the presence of mitotic cohesins SA and Nipped-B, this pathway has only a minor role in meiotic sister-centromere cohesion and is primarily required for homolog interactions. C(2)M is continuously incorporated into pachytene chromosomes even though SC assembly is complete. In contrast, the second complex, which depends on meiosis-specific proteins SOLO, SUNN, and ORD is required for sister-chromatid cohesion, localizes to the centromeres and is not incorporated during prophase. Our results show that the two cohesin complexes have unique functions and are regulated differently. Multiple cohesin complexes may provide the diversity of activities required by the meiotic cell. For example, a dynamic complex may allow the chromosomes to regulate meiotic recombination, and a stable complex may be required for sister-chromatid cohesion.

Isolated and Dynamical Horizons and Their Applications.

Over the past three decades, black holes have played an important role in quantum gravity, mathematical physics, numerical relativity and gravitational wave phenomenology. However, conceptual settings and mathematical models used to discuss them have varied considerably from one area to another. Over the last five years a new, quasi-local framework was introduced to analyze diverse facets of black holes in a unified manner. In this framework, evolving black holes are modelled by dynamical horizons and black holes in equilibrium by isolated horizons. We review basic properties of these horizons and summarize applications to mathematical physics, numerical relativity, and quantum gravity. This paradigm has led to significant generalizations of several results in black hole physics. Specifically, it has introduced a more physical setting for black hole thermodynamics and for black hole entropy calculations in quantum gravity, suggested a phenomenological model for hairy black holes, provided novel techniques to extract physics from numerical simulations, and led to new laws governing the dynamics of black holes in exact general relativity.

Sisters unbound is required for meiotic centromeric cohesion in Drosophila melanogaster.

Regular meiotic chromosome segregation requires sister centromeres to mono-orient (orient to the same pole) during the first meiotic division (meiosis I) when homologous chromosomes segregate, and to bi-orient (orient to opposite poles) during the second meiotic division (meiosis II) when sister chromatids segregate. Both orientation patterns require cohesion between sister centromeres, which is established during meiotic DNA replication and persists until anaphase of meiosis II. Meiotic cohesion is mediated by a conserved four-protein complex called cohesin that includes two structural maintenance of chromosomes (SMC) subunits (SMC1 and SMC3) and two non-SMC subunits. In Drosophila melanogaster, however, the meiotic cohesion apparatus has not been fully characterized and the non-SMC subunits have not been identified. We have identified a novel Drosophila gene called sisters unbound (sunn), which is required for stable sister chromatid cohesion throughout meiosis. sunn mutations disrupt centromere cohesion during prophase I and cause high frequencies of non-disjunction (NDJ) at both meiotic divisions in both sexes. SUNN co-localizes at centromeres with the cohesion proteins SMC1 and SOLO in both sexes and is necessary for the recruitment of both proteins to centromeres. Although SUNN lacks sequence homology to cohesins, bioinformatic analysis indicates that SUNN may be a structural homolog of the non-SMC cohesin subunit stromalin (SA), suggesting that SUNN may serve as a meiosis-specific cohesin subunit. In conclusion, our data show that SUNN is an essential meiosis-specific Drosophila cohesion protein.

Dynamical horizons: energy, angular momentum, fluxes, and balance laws.

Dynamical horizons are considered in full, nonlinear general relativity. Expressions of fluxes of energy and angular momentum carried by gravitational waves across these horizons are obtained. Fluxes are local, the energy flux is positive, and change in the horizon area is related to these fluxes. The flux formulas also give rise to balance laws analogous to the ones obtained by Bondi and Sachs at null infinity and provide generalizations of the first and second laws of black-hole mechanics.