Skull Architecture Research Articles

Variation in the cranial osteology of the amphisbaenian genus Zygaspis based on high-resolution x-ray computed tomography.

Amphisbaenians are a specialized fossorial group of reptiles, having developed head-first burrowing, a specialized skull architecture, and an elongated body. This group is generally small-bodied, with some species possessing skulls only a few millimeters long. In this study, we used high-resolution x-ray computed tomography to compare the skulls of 15 specimens from seven of the eight species in the amphisbaenian genus Zygaspis (Zygaspis dolichomenta, Zygaspis ferox, Zygaspis quadrifrons, Zygaspis kafuensis, Zygaspis nigra, Zygaspis vandami, and Zygaspis violacea). Both interspecific and intraspecific variation, including asymmetry, is observed among the cranial bones of the specimens. There are unique morphological features on some cranial bones, including the premaxilla and ectopterygoid of Z. quadrifrons, the pterygoid and vomer of Z. kafuensis, and the extracolumella of Z. nigra. Sexual dimorphism has been previously reported for the species Z. quadrifrons and is observed here as well.

Convergence, divergence, and macroevolutionary constraint as revealed by anatomical network analysis of the squamate skull, with an emphasis on snakes

Traditionally considered the earliest-diverging group of snakes, scolecophidians are central to major evolutionary paradigms regarding squamate feeding mechanisms and the ecological origins of snakes. However, quantitative analyses of these phenomena remain scarce. Herein, we therefore assess skull modularity in squamates via anatomical network analysis, focusing on the interplay between ‘microstomy’ (small-gaped feeding), fossoriality, and miniaturization in scolecophidians. Our analyses reveal distinctive patterns of jaw connectivity across purported ‘microstomatans’, thus supporting a more complex scenario of jaw evolution than traditionally portrayed. We also find that fossoriality and miniaturization each define a similar region of topospace (i.e., connectivity-based morphospace), with their combined influence imposing further evolutionary constraint on skull architecture. These results ultimately indicate convergence among scolecophidians, refuting widespread perspectives of these snakes as fundamentally plesiomorphic and morphologically homogeneous. This network-based examination of skull modularity—the first of its kind for snakes, and one of the first to analyze squamates—thus provides key insights into macroevolutionary trends among squamates, with particular implications for snake origins and evolution.

A Triassic stem lepidosaur illuminates the origin of lizard-like reptiles.

The early evolution of diapsid reptiles is marked by a deep contrast between our knowledge of the origin and early evolution of archosauromorphs (crocodiles, avian and non-avian dinosaurs) to that of lepidosauromorphs (squamates (lizards, snakes) and sphenodontians (tuataras)). Whereas the former include hundreds of fossil species across various lineages during the Triassic period1, the latter are represented by an extremely patchy early fossil record comprising only a handful of fragmentary fossils, most of which have uncertain phylogenetic affinities and are confined to Europe1-3. Here we report the discovery of a three-dimensionally preserved reptile skull, assigned as Taytalura alcoberi gen. et sp. nov., from the Late Triassic epoch of Argentina that is robustly inferred phylogenetically as the earliest evolving lepidosauromorph, using various data types and optimality criteria. Micro-computed tomography scans of this skull reveal details about the origin of the lepidosaurian skull from early diapsids, suggesting thatseveral traits traditionally associated with sphenodontians in fact originated much earlier in lepidosauromorph evolution. Taytalura suggests that the strongly evolutionarily conserved skull architecture of sphenodontians represents the plesiomorphic condition for all lepidosaurs, that stem and crown lepidosaurs were contemporaries for at least ten million years during the Triassic, and that early lepidosauromorphs had a much broader geographical distribution than has previously been thought.

Ontogenetic and in silico models of spatial-packing in the hypermuscular mouse skull.

Networks linking single genes to multiple phenotypic outcomes can be founded on local anatomical interactions as well as on systemic factors like biochemical products. Here we explore the effects of such interactions by investigating the competing spatial demands of brain and masticatory muscle growth within the hypermuscular myostatin-deficient mouse model and in computational simulations. Mice that lacked both copies of the myostatin gene (-/-) and display gross hypermuscularity, and control mice that had both copies of the myostatin gene (+/+) were sampled at 1, 7, 14 and 28 postnatal days. A total of 48 mice were imaged with standard as well as contrast-enhanced microCT. Size metrics and landmark configurations were collected from the image data and were analysed alongside in silico models of tissue expansion. Findings revealed that: masseter muscle volume was smaller in -/- mice at day 1 but became, and remained thereafter, larger by 7days; -/- endocranial volumes begin and remained smaller; -/- enlargement of the masticatory muscles was associated with caudolateral displacement of the calvarium, lateral displacement of the zygomatic arches, and slight dorsal deflection of the face and basicranium. Simulations revealed basicranial retroflexion (flattening) and dorsal deflection of the face associated with muscle expansion and abrogative covariations of basicranial flexion and ventral facial deflection associated with endocranial expansion. Our findings support the spatial-packing theory and highlight the importance of understanding the harmony of competing spatial demands that can shape and maintain mammalian skull architecture during ontogeny.

Feeding biomechanics suggests progressive correlation of skull architecture and neck evolution in turtles

The origin of turtles is one of the most long-lasting debates in evolutionary research. During their evolution, a series of modifications changed their relatively kinetic and anapsid skull into an elongated akinetic structure with a unique pulley system redirecting jaw adductor musculature. These modifications were thought to be strongly correlated to functional adaptations, especially to bite performance. We conducted a series of Finite Element Analyses (FEAs) of several species, including that of the oldest fully shelled, Triassic stem-turtle Proganochelys, to evaluate the role of force distribution and to test existing hypotheses on the evolution of turtle skull architecture. We found no support for a relation between the akinetic nature of the skull or the trochlear mechanisms with increased bite forces. Yet, the FEAs show that those modifications changed the skull architecture into an optimized structure, more resistant to higher loads while allowing material reduction on specific regions. We propose that the skull of modern turtles is the result of a complex process of progressive correlation between their heads and highly flexible necks, initiated by the origin of the shell.

Osteology of the archosauromorph Teyujagua paradoxa and the early evolution of the archosauriform skull

Abstract Archosauriformes are a major group of fossil and living reptiles that include the crown group Archosauria (birds, crocodilians and their extinct relatives) and closely related taxa. Archosauriformes are characterized by a highly diagnostic skull architecture, which is linked to the predatory habits of their early representatives, and the development of extensive cranial pneumaticity associated with the nasal capsule. The evolution of the archosauriform skull from the more plesiomorphic configuration present ancestrally in the broader clade Archosauromorpha was, until recently, elusive. This began to change with the discovery and description of Teyujagua paradoxa, an early archosauromorph from the Lower Triassic Sanga do Cabral Formation of Brazil. Here, we provide a detailed osteological description of the holotype and, thus far, only known specimen of T. paradoxa. In addition to providing new details of the anatomy of T. paradoxa, our study also reveals an early development of skull pneumaticity prior to the emergence of the antorbital fenestra. We use these new data to discuss the evolution of antorbital openings within Archosauriformes. Reappraisal of the phylogenetic position of T. paradoxa supports previous hypotheses of a close relationship with Archosauriformes. The data presented here provide new insights into character evolution during the origin of the archosauriform skull.

Unique skull network complexity of Tyrannosaurus rex among land vertebrates

Like other diapsids, Tyrannosaurus rex has two openings in the temporal skull region. In addition, like in other dinosaurs, its snout and lower jaw show large cranial fenestrae. In T. rex, they are thought to decrease skull weight, because, unlike most other amniotes, the skull proportion is immense compared to the body. Understanding morphofunctional complexity of this impressive skull architecture requires a broad scale phylogenetic comparison with skull types different to that of dinosaurs with fundamentally diverging cranial regionalization. Extant fully terrestrial vertebrates (amniotes) provide the best opportunities in that regard, as their skull performance is known from life. We apply for the first time anatomical network analysis to study skull bone integration and modular constructions in tyrannosaur and compare it with five representatives of the major amniote groups in order to get an understanding of the general patterns of amniote skull modularity. Our results reveal that the tyrannosaur has the most modular skull organization among the amniotes included in our study, with an unexpected separation of the snout in upper and lower sub-modules and the presence of a lower adductor chamber module. Independent pathways of bone reduction in opossum and chicken resulted in different degrees of cranial complexity with chicken having a typical sauropsidian pattern. The akinetic skull of opossum, alligator, and leatherback turtle evolved in independent ways mirrored in different patterns of skull modularity. Kinetic forms also show great diversity in modularity. The complex tyrannosaur skull modularity likely represents a refined mosaic of phylogenetic and ecological factors with food processing being probably most important for shaping its skull architecture. Mode of food processing primarily shaped skull integration among amniotes, however, phylogenetic patterns of skull integration are low in our sampling. Our general conclusions on amniote skull integrity are obviously preliminary and should be tested in subsequent studies. As such, this study provides a framework for future research focusing on the evolution of modularity on lower taxonomic levels.

Dinosaur ossification centres in embryonic birds uncover developmental evolution of the skull.

Radical transformation of the skull characterizes bird evolution. An increase in the relative size of the brain and eyes was presumably related to the loss of two bones surrounding the eye, the prefrontal and postorbital. We report that ossification centres of the prefrontal and postorbital are still formed in bird embryos, which then fuse seamlessly to the developing nasal and frontal bones, respectively, becoming undetectable in the adult. The presence of a dinosaur-like ossification pattern in bird embryos is more than a trace of their evolutionary past: we show how persistent modularity of ossification centres has allowed for evolutionary re-organization of skull architecture in evolution. Our findings also demonstrate that enigmatic mesodermal cells forming the posterior region of the avian frontal correspond to the ossification centre of the postorbital, not the parietal, and link its failure to develop into an adult bone to its incorporation into the expanded braincase of birds.

Custom made orthotic device for maintaining skull architecture during the postoperative period in infants undergoing craniosynostosis surgery

AimTo fabricate a cost effective, indigenous and simple orthotics helmet for post-operative cranial molding in patients with craniosynostosis surgery. MethodsWe present a case of 15 month old infant with secondary cranial vault deformity. Cranial vault remodeling surgery involving the posterior skull was planned and executed to increase the posterior gap, so that brain growth would be facilitated towards this empty space. Materials such as thermoplastic sponge, thermoplastic ionomer resin sheet, soft sponge and Velcro straps are used to fabricate a cranial orthotics helmet. ResultsWe have successfully used the above materials to fabricate the orthotics helmet for post-operative cranial molding. ConclusionThe technique described in this article is simple and cost effective. It can be custom made according to the demands of the surgical technique and the type of synostosis. It favors an individualistic prognosis, and proves worthwhile as every synostosis requires a unique treatment plan. It is an excellent adjuvant to craniosynostosis remodeling surgery.

The importance of accurate muscle modelling for biomechanical analyses: a case study with a lizard skull

Computer-based simulation techniques such as multi-body dynamics analysis are becoming increasingly popular in the field of skull mechanics. Multi-body models can be used for studying the relationships between skull architecture, muscle morphology and feeding performance. However, to be confident in the modelling results, models need to be validated against experimental data, and the effects of uncertainties or inaccuracies in the chosen model attributes need to be assessed with sensitivity analyses. Here, we compare the bite forces predicted by a multi-body model of a lizard (Tupinambis merianae) with in vivo measurements, using anatomical data collected from the same specimen. This subject-specific model predicts bite forces that are very close to the in vivo measurements and also shows a consistent increase in bite force as the bite position is moved posteriorly on the jaw. However, the model is very sensitive to changes in muscle attributes such as fibre length, intrinsic muscle strength and force orientation, with bite force predictions varying considerably when these three variables are altered. We conclude that accurate muscle measurements are crucial to building realistic multi-body models and that subject-specific data should be used whenever possible.

Feeding-related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet

Ősi, A. 2011: Feeding-related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet. Lethaia, Vol. 44, pp. 136–152. A comparative study of various feeding-related features in basal pterosaurs reveals a significant change in feeding strategies during the early evolutionary history of the group. These features are related to the skull architecture (e.g. quadrate morphology and orientation, jaw joint), dentition (e.g. crown morphology, wear patterns), reconstructed adductor musculature and post-cranium. The most basal pterosaurs (Preondactylus, dimorphodontids and anurognathids) were small-bodied animals with a wingspan no greater than 1.5 m, a relatively short, lightly constructed skull, straight mandibles with a large gape, sharply pointed teeth and well-developed external adductors. The absence of extended tooth wear excludes complex oral food processing and indicates that jaw closure was simply orthal. Features of these basal-most forms indicate a predominantly insectivorous diet. Among stratigraphically older but more derived forms (Eudimorphodon, Carniadactylus, Caviramus) complex, multicuspid teeth allowed the consumption of a wider variety of prey via a more effective form of food processing. This is supported by heavy dental wear in all forms with multicuspid teeth. Typical piscivorous forms occurred no earlier than the Early Jurassic, and are characterized by widely spaced, enlarged procumbent teeth forming a fish grab and an anteriorly inclined quadrate that permitted only a relatively small gape. In addition, the skull became more elongate and body size increased. Besides the dominance of piscivory, dental morphology and the scarcity of tooth wear reflect accidental dental occlusion that could have been caused by the capturing or seasonal consumption of harder food items. □Basal pterosaurs, heterodonty, dental wear, insectivory, piscivory.

Do agility and skull architecture influence the geometry of the mammalian vestibulo‐ocular reflex?

The spatial arrangement of the semicircular canals and extraocular muscles of the eye has been of considerable interest, particularly to researchers working on adaptations of the vestibulo-ocular reflex. Here we offer the first, extensive comparative analysis of the spatial relationships between each extraocular muscle and the canal providing its primary excitatory stimulus. The sample consisted of 113 specimens, representing 51 extant mammalian species. Hypotheses tested included that variations in the spatial alignments are linked with differences of skull morphology and with differences of agility during locomotion. Internal morphologies were visualized with magnetic resonance imaging and were measured with landmark-based vectors and planes. Values for body mass and agility were taken from the existing literature. Data were investigated for trends and associations with standard bivariate and multivariate statistical methods as well as with phylogenetically adjusted bivariate methods. The findings clearly show that species differences in the alignment of each extraocular muscle relative to the canal providing its primary excitatory stimulus are closely associated with changes of orbit morphology. The results also indicate that the actions of the oblique muscles interchange with those of the superior and inferior recti muscles when comparing lateral-eyed (rabbit) with frontal-eyed species (cat). There was only weak evidence to support the notion that canal-muscle alignments differ significantly among species according to how agile they are. The results suggest that semicircular canal morphology is arranged primarily for detecting head movements and then secondarily, if at all, for diminishing the burden of transforming vestibulo-ocular reflex signals in the most agile species.

New occurrence of the genus Dakosaurus (Reptilia, Thalattosuchia) in the Upper Jurassic of north-eastern Mexico, with comments upon skull architecture of Dakosaurus and Geosaurus

New occurrence of the genus Dakosaurus (Reptilia, Thalattosuchia) in the Upper Jurassic of north-eastern Mexico, with comments upon skull architecture of Dakosaurus and Geosaurus

Craniofacial growth in fetal Tarsius bancanus: brains, eyes and nasal septa.

The tarsier skull has been of particular interest in studies of primate taxonomy and functional morphology for several decades. Despite this, there remains no comprehensive data on how the tarsier skull develops, especially in relation to the soft-tissues of the head. Here we have documented for the first time fetal development of the skull and brain as well as the nasal septum and eyes in T. bancanus. We have also tested for the possible influence of these tissues in shaping skull architecture. Nineteen post-mortem specimens were imaged using high-resolution magnetic resonance imaging and magnetic resonance microscopy. Landmarks and volume data were collected and analysed. Findings demonstrated massive increases of brain size and eye size as well as flattening of the midline cranial base, facial projection and orbital margin frontation. Little evidence was found to support the notion that growth of the brain or nasal septum physically drives the observed changes of the skull. However, increases in the size of the eyes relative to skull size were associated with orbital margin frontation. With the possible exception of the results for eye size, the findings indicate that rather than forcing change the soft-tissues form a framework that physically constrains the morphogenetic template of the skeletal elements. This suggests, for example, that the degree of cranial base angulation seen in adulthood is not directly determined by brain expansion bending the basicranium, but by brain enlargement limiting the extent of cranial base flattening (retroflexion) in the fetus.

Extensive enlargement of the maxillary sinus inAlouatta caraya(mammalia, primates, cebidae): An allometric approach to skull pneumatization in Atelinae

In contrast to the paranasal sinuses of Old World monkeys and hominoids, little information is available about the paranasal sinuses of New World monkeys. Because this information is crucial in order to draw further conclusions about the evolution and biological role of skull pneumatization, this study investigates the morphology of the paranasal sinuses in adult black-and-gold howler monkeys (Alouatta caraya). Volumes of the paranasal sinuses were calculated using computer software (SURFdriver or Allegro) from serial coronal CT scans of 20 skulls of both sexes. Skull pneumatization in A. caraya is more complex than in other higher primates. In both sexes, the maxillary sinus (MS) is the only pneumatic cavity and enlarges regularly into neighboring bones such as the frontal bone and the basisphenoid. The resulting pansinus is often partitioned by several vertical septa. As in most external cranial dimensions, mean MS volume of A. caraya (male 4.08 cm(3); female 2.00 cm(3)) shows significant sexual dimorphism. Reduced major axis regression analysis between MS volume and different cranial dimensions for A. caraya (and for available data from other platyrrhines) suggests a distinct association for this group, with Alouatta having one of the largest pneumatic cavities. The combination of this unusual expansion of the MS of Alouatta and the occurrence of distinct septa within the sinus may be a consequence of the distinct skull architecture of Alouatta.

The correlation between skull fractures and intracranial lesions due to traffic accidents.

In this study, it was aimed to investigate the relationship between skull fractures and intracranial lesions following head injury. For this purpose, 500 cases, which were referred to the Third Committee of Council of Forensic Medicine in Istanbul due to traffic accidents by the courts of laws between 1998 and 2000, were examined retrospectively. They were categorized in 3 groups based on findings of their cranium x-rays and brain tomographies. 1- The cases who have fractures in skull bones with brain lesions 2- The cases who have fractures in skull bones with no brain lesions 3- The cases who have brain lesions with no skull fractures. They were examined in detail according to age, sex, localization of skull fractures and brain lesions, and if surgery was applied or not. Of the cases, 152 (30.4%) had only linear fractures, 69 (13.8%) had depressed fractures, 92 (18.4%) had linear fractures plus intracranial lesions, 49 (9.8%) had depressed fractures plus intracranial lesions and 138 (27.6%) had only intracranial lesions. The rate of intracranial lesion among the cases with the skull fracture was 38.9% (141/362), while the rate of skull fracture among the cases with the intracranial lesion was 50.3% (141/279) (P < 0.001). Male to female ratios were 2.4/1 for linear fractures, 5.2/1 for depressed fractures, and 3.5/1 for intracranial lesions. Linear fractures were more frequent among females whereas depressed fractures were often among males (chi2: 9.68, df: 4, p: 0.046). The mean age was 26.3. The rate of depressed fractures was higher the age groups of 0-30 years. (chi2: 16.28, df: 4, p: 0.003). Depressed fractures in the regions of frontal and parietal and, linear fracture in the regions of temporal and occipital were found at higher rates (P < 0.001). In conclusion, we reviewed skull fractures and/or intracranial lesions due to traffic accidents, and found depressed fractures to be more common among males whereas linear fractures to be more common among females and young males. In the male, the skull architecture is thicker and stronger than females and young males. We can state that presence of skull fractures lowers the incidence of intracranial lesions by lowering the intracranial pressure.

A redescription of the skull of Euoplocephalus tutus (Archosauria: Ornithischia): a foundation for comparative and systematic studies of ankylosaurian dinosaurs

Euoplocephalus tutusLambe (1902) from the Late Cretaceous of North America, was the first ankylosaurian dinosaur known from significant cranial material. Previous descriptions of this and other members of the Ankylosauria have been constrained by a paucity of material and an extremely apomorphic skull architecture, including the pervasive development of an embossing osseous ornamentation, and the absence of traditional morphological landmarks. A relative abundance of more recently collected and prepared cranial material attributable to Euoplocephalus enables a reappraisal of this taxon (including the type specimen), and permits it to be employed as a morphological representative of the clade. In recognition of previous difficulties encountered due to peculiarities of ankylosaurian anatomy, a fresh descriptive approach is necessitated. Herein, the skull is subdivided into five mutually exclusive topographic regions, within which individual elements are assigned with the assistance of outgroup comparison. Euoplocephalus is characterized by a distinctive pattern of cranial sculpturing across the preorbital area, relatively small, variably fluted teeth lacking a cingulum, a modified palpebral and a shallow nasal vestibule. Among ankylosaurine ankylosaurs, Euoplocephalus is unique in having medially inflected maxillary tooth rows. Osteological evaluation of the type skull of Anodontosaurus lambeiSternberg, 1929 supports its placement into synonymy with Euoplocephalus.

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best-known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well-developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda.

Pneumatized Spaces, Sinuses and Spongy Bones in the Skulls of Primates

The earliest attempts to understand the "pneumatized spaces" in the skulls of primates in general were focussed on the hollow spaces and the epithelium which covers their surfaces. More recent approaches consider the sinuses as a means to optimise skull architecture. Still, many attempts to get hold of the meaning of the intriguing pneumatized spaces circle around the air filled volumes they enclose. Here, we would like to reverse the approach and focus our biomechanic interpretation on the walls surrounding the big, empty, or at least not mechanically resistant spaces, and their mechanical properties. As a working hypothesis, we consider not only the walls of the more or less closed cavities, or sinuses, but also the braincase, the orbits, and the nasal channel as thin-walled shells of which we know that they can carry surprisingly large loads with a minimum of material. Details of the wall's profiles fit with this approach. From the same viewpoint, the bubble-like, air-filled cavernous systems in the ethmoid or temporal bones, and the marrow-filled spongy substance in the upper jaw are looked at as honeycomb-structures, which provide mechanical properties that are biologically advantageous and allow the saving of weight.

Cranial design and function in a large theropod dinosaur.

Finite element analysis (FEA) is used by industrial designers and biomechanicists to estimate the performance of engineered structures or human skeletal and soft tissues subjected to varying regimes of stress and strain. FEA is rarely applied to problems of biomechanical design in animals, despite its potential to inform structure-function analysis. Non-invasive techniques such as computed tomography scans can be used to generate accurate three-dimensional images of structures, such as skulls, which can form the basis of an accurate finite element model. Here we have applied this technique to the long skull of the large carnivorous theropod dinosaur Allosaurus fragilis. We have generated the most geometrically complete and complex FEA model of the skull of any extinct or extant organism and used this to test its mechanical properties and examine, in a quantitative way, long-held hypotheses concerning overall shape and function. The combination of a weak muscle-driven bite force, a very 'light' and 'open' skull architecture and unusually high cranial strength, suggests a very specific feeding behaviour for this animal. These results demonstrate simply the inherent potential of FEA for testing mechanical behaviour in fossils in ways that, until now, have been impossible.