Prof. Dr. John Nyakatura
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Biologische Adaptation: Eine integrative Studie zum Sprungverhalten und zur funktionellen Morphologie der Hinterextremitäten der Callitrichiden
Quelle ↗Förderer: DFG Sachbeihilfe Zeitraum: 12/2016 - 03/2023 Projektleitung: Prof. Dr. John Nyakatura, Kathleen Waak
EXC 1027: Bild Wissen Gestaltung. Ein Interdisziplinäres Labor
Quelle ↗Förderer: DFG Exzellenzinitiative Cluster Zeitraum: 11/2012 - 10/2017 Projektleitung: Prof. Dr. phil. Wolfgang Schäffner, Prof. Dr. Dr. h.c. Peter Fratzl, Prof. Dr. Horst Bredekamp
EXC 2025: Matters of Activity. Image Space Material
Quelle ↗Förderer: DFG Exzellenzstrategie Cluster Zeitraum: 01/2019 - 12/2025 Projektleitung: Prof. Dr. phil. Wolfgang Schäffner, Prof. Dr. Claudia Mareis
Functional Morphology of Stem Group Amniotes From the Lower Permian
Quelle ↗Förderer: Volkswagen Stiftung Zeitraum: 06/2015 - 12/2016 Projektleitung: Prof. Dr. John Nyakatura
Robotic Paleontology - Ein neuer Schlüssel zum Verständnis der frühen Evolution der Säugetiere
Quelle ↗Förderer: DFG sonstige Programme Zeitraum: 08/2026 - 07/2031 Projektleitung: Prof. Dr. John Nyakatura
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Publikationen25
Top 25 nach Zitationen — Quelle: OpenAlex (BAAI/bge-m3 embedded für Matching).
Nature · 223 Zitationen · DOI
American Journal of Physical Anthropology · 112 Zitationen · DOI
The influence of different substrate inclinations on gaits and metric gait parameters (relative forelimb and hind limb protraction, relative forelimb, and hind limb retraction, stride length, stance, and swing phase duration) of cotton-top tamarin locomotion was studied using high-speed video films and evaluated by descriptive and analytical statistical methods. As previously shown, lateral sequence gaits predominantly occurred on descending arboreal substrates (branchlike pole with a smaller diameter than the animal's body). Gait sequence patterns display significant dependency on substrate inclination. Cotton-top tamarins utilize lower diagonality values the more the substrate declines. This tendency leads to a greater use of lateral sequence gaits on steeply declined substrates. Conversely, these primates display the tendency to utilize higher diagonality values the more the substrate inclines leading to the predominant occurrence of diagonal sequence (DS) gaits. Duty factor index, extent of relative protraction, and relative retraction of both limb pairs as well as the relation of forelimb stance phase duration to hind limb stance phase duration is also correlated to the inclination of the substrate. Stride length and swing phase duration display no significant dependence on inclination, but are determined by the speed of the moving animal. The relevant duty factor is approximately constant at all inclinations. Integrating our results with results of other authors we propose a hypothesis for the functional relevance of a utilization of lateral sequence gaits in downward locomotion and DS gaits in upward locomotion. Our data support the notion of a wide ranging behavioral plasticity as a general primate locomotor characteristic.
Journal of Human Evolution · 108 Zitationen · DOI
Journal of Mammalian Evolution · 102 Zitationen · DOI
Frontiers in Zoology · 67 Zitationen · DOI
Most parameters were marked by an important intraspecific variability and by a size effect, which could, at least partly, be masking the functional signal. But for some parameters, the degree of anisotropy in particular, a clear functional distinction was recovered. Along with data on primates, our findings suggest that a trabecular architecture characterised by a greater degree of anisotropy is to be expected in species in which the relevant epiphyses withstand a restricted range of load directions. Trabecular architecture therefore is a promising research avenue for the reconstruction of lifestyles in extinct or cryptic species.
PLoS ONE · 64 Zitationen · DOI
Orobates pabsti, a basal diadectid from the lower Permian, is a key fossil for the understanding of early amniote evolution. Quantitative analysis of anatomical information suffers from fragmentation of fossil bones, plastic deformation due to diagenetic processes and fragile preservation within surrounding rock matrix, preventing further biomechanical investigation. Here we describe the steps taken to digitally reconstruct MNG 10181, the holotype specimen of Orobates pabsti, and subsequently use the digital reconstruction to assess body mass, position of the centre of mass in individual segments as well as the whole animal, and study joint mobility in the shoulder and hip joints. The shape of most fossil bone fragments could be recovered from micro-focus computed tomography scans. This also revealed structures that were hitherto hidden within the rock matrix. However, parts of the axial skeleton had to be modelled using relevant isolated bones from the same locality as templates. Based on the digital fossil, mass of MNG 10181 was estimated using a model of body shape that was varied within a plausible range to account for uncertainties of the dimension. In the mean estimate model the specimen had an estimated mass of circa 4 kg. Varying of the mass distribution amongst body segments further revealed that Orobates carried most of its weight on the hind limbs. Mostly unrestricted joint morphology further suggested that MNG 10181 was able to effectively generate propulsion with the pelvic limbs. The digital reconstruction is made available for future biomechanical studies.
Zoology · 62 Zitationen · DOI
Zoological Letters · 59 Zitationen · DOI
This study on sciuromorph trabeculae supplements the few non-primate studies on lifestyle-related functional adaptation of trabecular bone. We show that the architecture of the femoral head trabeculae in Sciuromorpha correlates with body mass and locomotor habits. Our findings provide a new basis for experimental research focused on functional significance of bone inner microstructure.
Proceedings of the Royal Society B Biological Sciences · 59 Zitationen · DOI
In contrast to the upright trunk in humans, trunk orientation in most birds is almost horizontal (pronograde). It is conceivable that the orientation of the heavy trunk strongly influences the dynamics of bipedal terrestrial locomotion. Here, we analyse for the first time the effects of a pronograde trunk orientation on leg function and stability during bipedal locomotion. For this, we first inferred the leg function and trunk control strategy applied by a generalized small bird during terrestrial locomotion by analysing synchronously recorded kinematic (three-dimensional X-ray videography) and kinetic (three-dimensional force measurement) quail locomotion data. Then, by simulating quail gaits using a simplistic bioinspired numerical model which made use of parameters obtained in in vivo experiments with real quail, we show that the observed asymmetric leg function (left-skewed ground reaction force and longer leg at touchdown than at lift-off) is necessary for pronograde steady-state locomotion. In addition, steady-state locomotion becomes stable for specific morphological parameters. For quail-like parameters, the most common stable solution is grounded running, a gait preferred by quail and most of the other small birds. We hypothesize that stability of bipedal locomotion is a functional demand that, depending on trunk orientation and centre of mass location, constrains basic hind limb morphology and function, such as leg length, leg stiffness and leg damping.
Journal of Experimental Biology · 55 Zitationen · DOI
Previous research has resulted in increasing insight into neuro-mechanical control strategies during perturbed locomotion. In contrast, more general analyses on simple model (template)-related parameters during avian terrestrial locomotion are still rare. Quail kinematic data obtained using X-ray videography combined with ground reaction force measurements were used as a basis to investigate how 'global' template and 'local' leg joint parameters in this small, predominantly terrestrial bird change with speed and gait. Globally, quail locomotion approximates a spring-like behavior in all investigated gaits. However, ground reaction forces are more vertically oriented, which may help to balance the trunk. At the joint level, practically all the spring-like work was found to occur in the intertarsal joint (ITJ). From walking to grounded running, the local stiffness of the ITJ decreases similarly to the reduction observed in global leg stiffness. Thus, in gaits without aerial phases the quails may modulate ITJ stiffness to regulate global leg stiffness, and therefore gait changes, to a significant degree. At higher speeds both global leg compression and stiffness are increased (the latter to values not significantly different to those obtained during walking). This enables the animals to shorten contact time and to generate aerial phases (running). However, we did not observe a change in the stiffness in the ITJ with a change of gait from grounded running to running. We hypothesize that a more extended leg at touch-down, controlled by the joint angles in the knee and ITJ, has an important influence in the leg stiffness adjustment process during running.
BMC Evolutionary Biology · 53 Zitationen · DOI
Although only future developmental analyses in non-model organisms, such as sloths, will yield direct evidence for the evolutionary mechanism responsible for the aberrant number of cervical vertebrae, our observations lend support to hypothesis 1 indicating that the number of modules is retained but their boundaries are displaced. Our approach based on quantified morphological differences also provides a reliable basis for further research including fossil taxa such as extinct 'ground sloths' in order to trace the pattern and the underlying genetic mechanisms in the evolution of the vertebral column in mammals.
Journal of Anatomy · 53 Zitationen · DOI
The reconstruction of a joint's maximum range of mobility (ROM) often is a first step when trying to understand the locomotion of fossil tetrapods. But previous studies suggest that the ROM of a joint is restricted by soft tissues surrounding the joint. To expand the limited informative value of ROM studies for the reconstruction of a fossil species' locomotor characteristics, it is moreover necessary to better understand the relationship of ex vivo ROM with the actual in vivo joint movement. To gain insight into the relationship between ex vivo mobility and in vivo movement, we systematically tested for the influence of soft tissues on joint ROM in the hip of the modern lizard Iguana iguana. Then, we compared the ex vivo mobility to in vivo kinematics of the hip joint in the same specimens using X-ray sequences of steady-state treadmill locomotion previously recorded. With stepwise removal of soft tissues and a repeated-measurement protocol, we show that soft tissues surrounding the hip joint considerably limit ROM, highlighting the problems when joint ROM is deduced from bare bones only. We found the integument to have the largest effect on the range of long-axis rotation, pro- and retraction. Importantly, during locomotion the iguana used only a fragment of the ROM that was measured in our least restrictive dissection situation (i.e. pelvis and femur only conjoined by ligaments), demonstrating the discrepancy between hip joint ROM and actual in vivo movement. Our study emphasizes the necessity for caution when attempting to reconstruct joint ROM or even locomotor kinematics from fossil bones only, as actual in vivo movement cannot be deduced directly from any condition of cadaver mobility in Iguana and likely in other tetrapods.
Evolutionary Biology · 52 Zitationen · DOI
Inertial Tail Effects during Righting of Squirrels in Unexpected Falls: From Behavior to Robotics
2021Integrative and Comparative Biology · 50 Zitationen · DOI
Arboreal mammals navigate a highly three dimensional and discontinuous habitat. Among arboreal mammals, squirrels demonstrate impressive agility. In a recent "viral" YouTube video, unsuspecting squirrels were mechanically catapulted off of a track, inducing an initially uncontrolled rotation of the body. Interestingly, they skillfully stabilized themselves using tail motion, which ultimately allowed the squirrels to land successfully. Here we analyze the mechanism by which the squirrels recover from large body angular rates. We analyzed from the video that squirrels first use their tail to help stabilizing their head to visually fix a landing site. Then the tail starts to rotate to help stabilizing the body, preparing themselves for landing. To analyze further the mechanism of this tail use during mid-air, we built a multibody squirrel model and showed the righting strategy based on body inertia moment changes and active angular momentum transfer between axes. To validate the hypothesized strategy, we made a squirrel-like robot and demonstrated a fall-stabilizing experiment. Our results demonstrate that a squirrel's long tail, despite comprising just 3% of body mass, can inertially stabilize a rapidly rotating body. This research contributes to better understanding the importance of long tails for righting mechanisms in animals living in complex environments such as trees.
Journal of Anatomy · 49 Zitationen · DOI
Recent phylogenetic analyses imply a diphyly of tree sloths and a convergent evolution of their obligatory suspensory locomotion. In mammals the extrinsic shoulder musculature forms a 'muscular sling' to support the trunk in quadrupedal postures. In addition, the extrinsic pectoral muscles are responsible for moving the proximal forelimb elements during locomotion. Due to the inverse orientation of the body in regard to the gravitational force, the muscular sling as configured as in pronograde mammals is unsuited to suspend the weight of the thorax in sloths. We here review the muscular topography of the shoulder in Choloepus didactylus and Bradypus variegatus in the light of presumably convergent evolution to adapt to the altered functional demands of the inverse orientation of the body. In addition, we venture to deduce the effect of the shoulder musculature of C. didactylus during locomotion based on previously published 3D kinematic data. Finally, we assess likely convergences in the muscular topography of both extant sloth lineages to test the hypothesis that convergent evolution is reflected by differing morphological solutions to the same functional demands posed by the suspensory posture. Muscular topography of the shoulder in C. didactylus is altered from the plesiomorphic condition of pronograde mammals, whereas the shoulder in B. variegatus more closely resembles the general pattern. Overall kinematics as well as the muscles suitable for pro- and retraction of the forelimb were found to be largely comparable to pronograde mammals in C. didactylus. We conclude that most of the peculiar topography of extrinsic forelimb musculature can be attributed to the inverse orientation of the body. These characteristics are often similar in both genera, but we also identified different morphological solutions that evolved to satisfy the new functional demands and are indicative of convergent evolution. We suggest that the shared phylogenetic heritage canalized the spectrum of possible solutions to new functional demands, and digging adaptations of early xenarthrans posed morphological constraints that resulted in similar suspensory postures. The data of this study, including muscle maps, will be helpful to infer locomotor characteristics of fossil sloths.
Journal of Experimental Biology · 49 Zitationen · DOI
Given the importance of thoraco-lumbar spine movements in the locomotion of mammals, it is surprising that in vivo three-dimensional (3-D) data on the intervertebral movement of the mammalian thoraco-lumbar vertebral column during symmetrical gaits is limited to horses and dogs. To test whether kinematic patterns similar to those published for these cursorial species are also present during a contrasting mode of quadrupedalism, we quantified thoraco-lumbar intervertebral movements, the resulting pelvic displacements and relative femoral movements during the trot-like steady-state suspensory quadrupedal locomotion of the two-toed sloth (Xenarthra, Choloepus didactylus). Scientific rotoscoping, a new, non-invasive approach that combines synchronous biplanar high speed X-ray videos and the reconstruction of skeletal elements from computed tomography bone scans, was used to quantify 3-D kinematics. An analysis of vertebral anatomy and epaxial muscle topography suggests that the thoraco-lumbar spine of sloths is well suited to producing lateral bending and long-axis rotation, but limits powerful sagittal extension. Sloths exhibit complex 3-D movements in the thoraco-lumbar spine that are comparable to those observed in other arboreal quadrupedal mammals. Monophasic lateral bending and long-axis rotation, biphasic sagittal bending and maximal amplitude of sagittal bending at the lumbo-sacral joint were also found in other quadruped mammals and may represent general aspects of mammalian symmetric gaits. Maximal amplitude of lateral bending and long-axis rotation vary in regard to the vertebral level. It is suggested that a cranio-caudal pattern of angular deflections of the spine results from the out-of-phase movement of diagonal forelimbs and hindlimbs in other walking gaits, because it is not evident in the trot-like locomotion analyzed here. The analysis also illustrates the difficulties that arise when lumbar movement is deduced from intervertebral joint morphology alone.
PLoS ONE · 48 Zitationen · DOI
A statistical analysis was used to evaluate the regionalization of the cervical spine in the barn owl. While earlier measurements have shown that there appear to be three regions of flexibility of the neck, our indicators suggest 3-7 regions. These many regions allow a high degree of flexibility, potentially facilitating the large head turns that barn owls are able to make. The cervical vertebral series of other species should also be investigated using statistical criteria to further characterize morphology and the potential movements associated with it.
Current Biology · 45 Zitationen · DOI
The elephant trunk operates as a muscular hydrostat<sup>1</sup><sup>,</sup><sup>2</sup> and is actuated by the most complex musculature known in animals.<sup>3</sup><sup>,</sup><sup>4</sup> Because the number of trunk muscles is unclear,<sup>5</sup> we performed dense reconstructions of trunk muscle fascicles, elementary muscle units, from microCT scans of an Asian baby elephant trunk. Muscle architecture changes markedly across the trunk. Trunk tip and finger consist of about 8,000 extraordinarily filigree fascicles. The dexterous finger consists exclusively of microscopic radial fascicles pointing to a role of muscle miniaturization in elephant dexterity. Radial fascicles also predominate (at 82% volume) the remainder of the trunk tip, and we wonder if radial muscle fascicles are of particular significance for fine motor control of the dexterous trunk tip. By volume, trunk-shaft muscles<sup>6</sup> comprise one-third of the numerous, small radial muscle fascicles; two-thirds of the three subtypes of large longitudinal fascicles (dorsal longitudinals, ventral outer obliques, and ventral inner obliques);<sup>7</sup><sup>,</sup><sup>8</sup><sup>,</sup><sup>9</sup> and a small fraction of transversal fascicles. Shaft musculature is laterally, but not radially, symmetric. A predominance of dorsal over ventral radial muscles and of ventral over dorsal longitudinal muscles may result in a larger ability of the shaft to extend dorsally than ventrally<sup>10</sup> and to bend inward rather than outward. There are around 90,000 trunk muscle fascicles. While primate hand control is based on fine control of contraction by the convergence of many motor neurons on a small set of relatively large muscles, evolution of elephant grasping has led to thousands of microscopic fascicles, which probably outnumber facial motor neurons.
Journal of Experimental Zoology Part A Ecological Genetics and Physiology · 44 Zitationen · DOI
Avian bipedalism is best studied in derived walking/running specialists. Here, we use kinematics and center of mass (CoM) mechanical energy patterns to investigate gait transitions of lapwings-migratory birds that forage on the ground, and therefore may need a trade-off between the functional demands of terrestrial locomotion and long distance flights. The animals ran on a treadmill while high-speed X-ray videos were recorded within the sustainable speed range. Instantaneous CoM mechanics were computed from integrating kinematics and body segment properties. Lapwings exhibit similar locomotor characteristics to specialized walking/running birds, but have less distinct gaits. At slow speeds no clear separation between vaulting (i.e., walking) and bouncing (i.e., running) energy patterns exists. Mechanical energy recovery of non-bouncing gaits correlates poorly with speed and suggests inefficient use of the inverted pendulum mechanism. Speed ranges of gaits overlap considerably, especially those of grounded running, a gait with CoM mechanics indicative of running but without an aerial phase, and aerial phase running, with no preferential gait at most speeds. Compliant limb morphology and grounded running in birds can be regarded as an evolutionary constraint, but lapwings effectively make use of advantages offered by this gait for a great fraction of their speed range. Thus, effective usage of grounded running during terrestrial locomotion is suggested generally to be a part of striding avian bipedalism-even in species not specialized in walking/running locomotion.
ACTA THERIOLOGICA · 39 Zitationen · DOI
Frontiers in Zoology · 39 Zitationen · DOI
The morphological characteristics of the scapula and the SCA allow maximal mobility of the forelimb to facilitate effective locomotion within a discontinuous habitat. These evolutionary changes associated with the adoption of the suspensory posture emphasized humeral influence on forelimb motion, but allowed the retention of the plesiomorphic 3D kinematic pattern.
Journal of Experimental Biology · 38 Zitationen · DOI
Comparative analyses of locomotion in tetrapods reveal two patterns of stride cycle variability. Tachymetabolic tetrapods (birds and mammals) have lower inter-cycle variation in stride duration than bradymetabolic tetrapods (amphibians, lizards, turtles and crocodilians). This pattern has been linked to the fact that birds and mammals share enlarged cerebella, relatively enlarged and heavily myelinated Ia afferents, and γ-motoneurons to their muscle spindles. Both tachymetabolic tetrapod lineages also possess an encapsulated Golgi tendon morphology, thought to provide more spatially precise information on muscle tension. The functional consequence of this derived Golgi tendon morphology has never been tested. We hypothesized that one advantage of precise information on muscle tension would be lower and more predictable limb bone stresses, achieved in tachymetabolic tetrapods by having less variable substrate reaction forces than bradymetabolic tetrapods. To test this hypothesis, we analyzed hindlimb substrate reaction forces during locomotion of 55 tetrapod species in a phylogenetic comparative framework. Variation in species means of limb loading magnitude and timing confirm that, for most of the variables analyzed, variance in hindlimb loading and timing is significantly lower in species with encapsulated versus unencapsulated Golgi tendon organs. These findings suggest that maintaining predictable limb loading provides a selective advantage for birds and mammals by allowing energy savings during locomotion, lower limb bone safety factors and quicker recovery from perturbations. The importance of variation in other biomechanical variables in explaining these patterns, such as posture, effective mechanical advantage and center-of-mass mechanics, remains to be clarified.
Journal of Anatomy · 37 Zitationen · DOI
Owls are known for their outstanding neck mobility: these birds can rotate their heads more than 270°. The anatomical basis of this extraordinary neck rotation ability is not well understood. We used X-ray fluoroscopy of living owls as well as forced neck rotations in dead specimens and computer tomographic (CT) reconstructions to study how the individual cervical joints contribute to head rotation in barn owls (Tyto furcata pratincola). The X-ray data showed the natural posture of the neck, and the reconstructions of the CT-scans provided the shapes of the individual vertebrae. Joint mobility was analyzed in a spherical coordinate system. The rotational capability was described as rotation about the yaw and roll axes. The analyses suggest a functional division of the cervical spine into several regions. Most importantly, an upper region shows high rolling and yawing capabilities. The mobility of the lower, more horizontally oriented joints of the cervical spine is restricted mainly to the roll axis. These rolling movements lead to lateral bending, effectively resulting in a side shift of the head compared with the trunk during large rotations. The joints in the middle of the cervical spine proved to contribute less to head rotation. The analysis of joint mobility demonstrated how owls might maximize horizontal head rotation by a specific and variable combination of yawing and rolling in functionally diverse regions of the neck.
Proceedings of the Royal Society B Biological Sciences · 37 Zitationen · DOI
Many ungulates show a conspicuous nodding motion of the head when walking. Until now, the functional significance of this behaviour remained unclear. Combining in vivo kinematics of quadrupedal mammals with a computer model, we show that the timing of vertical displacements of the head and neck is consistent with minimizing energy expenditure for carrying these body parts in an inverted pendulum walking gait. Varying the timing of head movements in the model resulted in increased metabolic cost estimate for carrying the head and neck of up to 63%. Oscillations of the head-neck unit result in weight force oscillations transmitted to the forelimbs. Advantageous timing increases the load in single support phases, in which redirecting the trajectory of the centre of mass (COM) is thought to be energetically inexpensive. During double support, in which-according to collision mechanics-directional changes of the impulse of the COM are expensive, the observed timing decreases the load. Because the head and neck comprise approximately 10% of body mass, the effect shown here should also affect the animals' overall energy expenditure. This mechanism, working analogously in high-tech backpacks for energy-saving load carriage, is widespread in ungulates, and provides insight into how animals economize locomotion.
Journal of Mammalian Evolution · 36 Zitationen · DOI
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- Prof. Dr. John Nyakatura
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