Gene flow from Corymbia hybrids in northern New South Wales

Some of the most productive taxa for forestry are interspecific F1 hybrids grown as exotics in the tropics and subtropics. Attributes of resilience, adaptability and vigour which engender the hybrids for wood production, may also exacerbate the risk they present from gene flow to native species gene pools or to local ecologies as weeds. To determine the biological and genetic factors that influence the extent of hybridisation, we examine the distribution and genealogy of wildlings surrounding plantings of locally-exotic Corymbia torelliana (Section Cadageria) near native C. henryi (Section Maculatae) in northern New South Wales. Our study showed pre-mating and pre- and post-zygotic barriers were incomplete, with in situ generation and natural establishment of both F1 hybrids (n = 3) and advanced generation hybrids under the disturbed conditions bordering native forest. As hybrids were located on alluvial flats exposed to frost, they also likely have an extended ecological range relative to native C. henryi. Despite the likely generation of large viable seed crops on F1 trees at the site over many years, establishment success and survival of advanced generation hybrids may be low, as only 5 immature and no mature advanced generation hybrids were identified. Propagation and genetic analysis of a seed crop from one F1 wildling showed early survival and vigour of seedlings in cultivation was high, and that at least for some F1 in some seasons, backcrossing to the recurrent native C. henryi parent is favoured (60%), whereas selfing (10%) and crossing with other F1 (30%) was less frequent. Transport of seed by stingless bees probably accounted for long distance dispersal from C. torelliana, but this mechanism does not appear to supplement gravity-dispersal of seed from the F1. Coupled with other evidence from studies of bee behaviour, controlled pollination in Corymbia sp., and long-term fitness in second generation eucalypt hybrids, we anticipate gene flow via pollen rather than seed will be the greater challenge for managing the risk of introgression of C. torelliana ancestry into native species from the planted F1 hybrid. If large sources of F1 pollen become available to compete with native pollen, gene flow will probably be frequent and hybrids may establish in disturbed conditions and in habitats beyond the ecological range of their native parent. Further study is needed to determine the degree to which outbreeding depression and poor survival inhibits on-going gene flow.

An evolutionary formalism for weak quantum measurements

Unitary evolution and projective measurement are fundamental axioms of quantum mechanics. Even though projective measurement yields one of the eigenstates of the measured operator as the outcome, there is no theory that predicts which eigenstate will be observed in which experimental run. There exists only an ensemble description, which predicts probabilities of various outcomes over many experimental runs. We propose a dynamical evolution equation for the projective collapse of the quantum state in individual experimental runs, which is consistent with the well-established framework of quantum mechanics. In case of gradual weak measurements, its predictions for ensemble evolution are different from those of the Born rule. It is an open question whether or not suitably designed experiments can observe this alternate evolution.

Global Self-Organization of the Cellular Metabolic Structure

Background: Over many years, it has been assumed that enzymes work either in an isolated way, or organized in small catalytic groups. Several studies performed using "metabolic networks models'' are helping to understand the degree of functional complexity that characterizes enzymatic dynamic systems. In a previous work, we used "dissipative metabolic networks'' (DMNs) to show that enzymes can present a self-organized global functional structure, in which several sets of enzymes are always in an active state, whereas the rest of molecular catalytic sets exhibit dynamics of on-off changing states. We suggested that this kind of global metabolic dynamics might be a genuine and universal functional configuration of the cellular metabolic structure, common to all living cells. Later, a different group has shown experimentally that this kind of functional structure does, indeed, exist in several microorganisms. Methodology/Principal Findings: Here we have analyzed around 2.500.000 different DMNs in order to investigate the underlying mechanism of this dynamic global configuration. The numerical analyses that we have performed show that this global configuration is an emergent property inherent to the cellular metabolic dynamics. Concretely, we have found that the existence of a high number of enzymatic subsystems belonging to the DMNs is the fundamental element for the spontaneous emergence of a functional reactive structure characterized by a metabolic core formed by several sets of enzymes always in an active state. Conclusions/Significance: This self-organized dynamic structure seems to be an intrinsic characteristic of metabolism, common to all living cellular organisms. To better understand cellular functionality, it will be crucial to structurally characterize these enzymatic self-organized global structures.

Characterization of diverse megathrust fault behavior related to seismic supercycles, Mentawai Islands, Sumatra

Long paleoseismic histories are necessary for understanding the full range of behavior of faults, as the most destructive events often have recurrence intervals longer than local recorded history. The Sunda megathrust, the interface along which the Australian plate subducts beneath Southeast Asia, provides an ideal natural laboratory for determining a detailed paleoseismic history over many seismic cycles. The outer-arc islands above the seismogenic portion of the megathrust cyclically rise and subside in response to processes on the underlying megathrust, providing uncommonly good illumination of megathrust behavior. Furthermore, the growth histories of coral microatolls, which record tectonic uplift and subsidence via relative sea level, can be used to investigate the detailed coseismic and interseismic deformation patterns. One particularly interesting area is the Mentawai segment of the megathrust, which has been shown to characteristically fail in a series of ruptures over decades, rather than a single end-to-end rupture. This behavior has been termed a seismic “supercycle.” Prior to the current rupture sequence, which began in 2007, the segment previously ruptured during the 14th century, the late 16th to late 17th century, and most recently during historical earthquakes in 1797 and 1833. In this study, we examine each of these previous supercycles in turn.

First, we expand upon previous analysis of the 1797–1833 rupture sequence with a comprehensive review of previously published coral microatoll data and the addition of a significant amount of new data. We present detailed maps of coseismic uplift during the two great earthquakes and of interseismic deformation during the periods 1755–1833 and 1950–1997 and models of the corresponding slip and coupling on the underlying megathrust. We derive magnitudes of Mw 8.7–9.0 for the two historical earthquakes, and determine that the 1797 earthquake fundamentally changed the state of coupling on the fault for decades afterward. We conclude that while major earthquakes generally do not involve rupture of the entire Mentawai segment, they undoubtedly influence the progression of subsequent ruptures, even beyond their own rupture area. This concept is of vital importance for monitoring and forecasting the progression of the modern rupture sequence.

Turning our attention to the 14th century, we present evidence of a shallow slip event in approximately A.D. 1314, which preceded the “conventional” megathrust rupture sequence. We calculate a suite of slip models, slightly deeper and/or larger than the 2010 Pagai Islands earthquake, that are consistent with the large amount of subsidence recorded at our study site. Sea-level records from older coral microatolls suggest that these events occur at least once every millennium, but likely far less frequently than their great downdip neighbors. The revelation that shallow slip events are important contributors to the seismic cycle of the Mentawai segment further complicates our understanding of this subduction megathrust and our assessment of the region’s exposure to seismic and tsunami hazards.

Finally, we present an outline of the complex intervening rupture sequence that took place in the 16th and 17th centuries, which involved at least five distinct uplift events. We conclude that each of the supercycles had unique features, and all of the types of fault behavior we observe are consistent with highly heterogeneous frictional properties of the megathrust beneath the south-central Mentawai Islands. We conclude that the heterogeneous distribution of asperities produces terminations and overlap zones between fault ruptures, resulting in the seismic “supercycle” phenomenon.

Toward Understanding Astrophysical Phenomena

Fast radio bursts (FRBs), a novel type of radio pulse, whose physics is not yet understood at all. Only a handful of FRBs had been detected when we started this project. Taking account of the scant observations, we put physical constraints on FRBs. We excluded proposals of a galactic origin for their extraordinarily high dispersion measures (DM), in particular stellar coronas and HII regions. Therefore our work supports an extragalactic origin for FRBs. We show that the resolved scattering tail of FRB 110220 is unlikely to be due to propagation through the intergalactic plasma. Instead the scattering is probably caused by the interstellar medium in the FRB's host galaxy, and indicates that this burst sits in the central region of that galaxy. Pulse durations of order $\ms$ constrain source sizes of FRBs implying enormous brightness temperatures and thus coherent emission. Electric fields near FRBs at cosmological distances would be so strong that they could accelerate free electrons from rest to relativistic energies in a single wave period. When we worked on FRBs, it was unclear whether they were genuine astronomical signals as distinct from `perytons', clearly terrestrial radio bursts, sharing some common properties with FRBs. Recently, in April 2015, astronomers discovered that perytons were emitted by microwave ovens. Radio chirps similar to FRBs were emitted when their doors opened while they were still heating. Evidence for the astronomical nature of FRBs has strengthened since our paper was published. Some bursts have been found to show linear and circular polarizations and Faraday rotation of the linear polarization has also been detected. I hope to resume working on FRBs in the near future. But after we completed our FRB paper, I decided to pause this project because of the lack of observational constraints.

The pulsar triple system, J0733+1715, has its orbital parameters fitted to high accuracy owing to the precise timing of the central $\ms$ pulsar. The two orbits are highly hierarchical, namely $P_{\mathrm{orb,1}}\ll P_{\mathrm{orb,2}}$, where 1 and 2 label the inner and outer white dwarf (WD) companions respectively. Moreover, their orbital planes almost coincide, providing a unique opportunity to study secular interaction associated purely with eccentricity beyond the solar system. Secular interaction only involves effect averaged over many orbits. Thus each companion can be represented by an elliptical wire with its mass distributed inversely proportional to its local orbital speed. Generally there exists a mutual torque, which vanishes only when their apsidal lines are parallel or anti-parallel. To maintain either mode, the eccentricity ratio, $e_1/e_2$, must be of the proper value, so that both apsidal lines precess together. For J0733+1715, $e_1\ll e_2$ for the parallel mode, while $e_1\gg e_2$ for the anti-parallel one. We show that the former precesses $\sim 10$ times slower than the latter. Currently the system is dominated by the parallel mode. Although only a little anti-parallel mode survives, both eccentricities especially $e_1$ oscillate on $\sim 10^3\yr$ timescale. Detectable changes would occur within $\sim 1\yr$. We demonstrate that the anti-parallel mode gets damped $\sim 10^4$ times faster than its parallel brother by any dissipative process diminishing $e_1$. If it is the tidal damping in the inner WD, we proceed to estimate its tidal quantity parameter ($Q$) to be $\sim 10^6$, which was poorly constrained by observations. However, tidal damping may also happen during the preceding low-mass X-ray binary (LMXB) phase or hydrogen thermal nuclear flashes. But, in both cases, the inner companion fills its Roche lobe and probably suffers mass/angular momentum loss, which might cause $e_1$ to grow rather than decay.

Several pairs of solar system satellites occupy mean motion resonances (MMRs). We divide these into two groups according to their proximity to exact resonance. Proximity is measured by the existence of a separatrix in phase space. MMRs between Io-Europa, Europa-Ganymede and Enceladus-Dione are too distant from exact resonance for a separatrix to appear. A separatrix is present only in the phase spaces of the Mimas-Tethys and Titan-Hyperion MMRs and their resonant arguments are the only ones to exhibit substantial librations. When a separatrix is present, tidal damping of eccentricity or inclination excites overstable librations that can lead to passage through resonance on the damping timescale. However, after investigation, we conclude that the librations in the Mimas-Tethys and Titan-Hyperion MMRs are fossils and do not result from overstability.

Rubble piles are common in the solar system. Monolithic elements touch their neighbors in small localized areas. Voids occupy a significant fraction of the volume. In a fluid-free environment, heat cannot conduct through voids; only radiation can transfer energy across them. We model the effective thermal conductivity of a rubble pile and show that it is proportional the square root of the pressure, $P$, for $P\leq \epsy^3\mu$ where $\epsy$ is the material's yield strain and $\mu$ its shear modulus. Our model provides an excellent fit to the depth dependence of the thermal conductivity in the top $140\,\mathrm{cm}$ of the lunar regolith. It also offers an explanation for the low thermal inertias of rocky asteroids and icy satellites. Lastly, we discuss how rubble piles slow down the cooling of small bodies such as asteroids.

Electromagnetic (EM) follow-up observations of gravitational wave (GW) events will help shed light on the nature of the sources, and more can be learned if the EM follow-ups can start as soon as the GW event becomes observable. In this paper, we propose a computationally efficient time-domain algorithm capable of detecting gravitational waves (GWs) from coalescing binaries of compact objects with nearly zero time delay. In case when the signal is strong enough, our algorithm also has the flexibility to trigger EM observation {\it before} the merger. The key to the efficiency of our algorithm arises from the use of chains of so-called Infinite Impulse Response (IIR) filters, which filter time-series data recursively. Computational cost is further reduced by a template interpolation technique that requires filtering to be done only for a much coarser template bank than otherwise required to sufficiently recover optimal signal-to-noise ratio. Towards future detectors with sensitivity extending to lower frequencies, our algorithm's computational cost is shown to increase rather insignificantly compared to the conventional time-domain correlation method. Moreover, at latencies of less than hundreds to thousands of seconds, this method is expected to be computationally more efficient than the straightforward frequency-domain method.

Polyphemus pediculus (L.) (Cladocera) as a possible indicator of water quality

Usually stenobiotic species are used as hydrobiological indicators of the degree of pollution in natural waters. Cladocera are eurybiotic organisms, therefore their role as specific indicators of the water quality is considered to be insignificant. However, considering new methods Cladocera at present are more and more often used as indicators of oligo- and mesosaprobic waters as well as of the presence of considerable amounts of easily degradable organic matter. Work over many years on the biology of Polyphemus pediculus, this striking representative of the order Cladocera, convinced the author of the possibility of using this species not only as an indicator of water purity but also for the estimation of the degree of water pollution as well as of water characteristics such as colour, turbidity, oxgen content and chemical composition. P. pediculus is one of the most common and abundant species of planktonic crustaceans in shallow waters of reservoirs, rivers, lakes, ponds and temporary water bodies.

Holistic fisheries management: combining macroecology, ecology, and evolutionary biology

Ecosystem-based management is one of many indispensable components of objective, holistic management of human impacts on nonhuman systems. By itself, however, ecosystem-based management carries the same risks we face with other forms of current management; holism requires more. Combining single-species and ecosystem approaches represents progress. However, it is now recognized that management also needs to be evosystem-based. In other words, management needs to account for all coevolutionary and evolutionary interactions among all species; otherwise we fall far short of holism. Fully holistic practices are quite distinct from the approaches to the management of fisheries that are applied today. In this paper, we show how macroecological patterns can guide management consistently, objectively, and holistically. We present one particular macroecological pattern with two applications. The first application is a case study of fisheries from the Baltic Sea involving historical data for two species; the second involves a sample of 44 species of primarily marine fish worldwide. In both cases we evaluate historical fishing rates and determine holistic/systemic sustainable single-species fishing rates to illustrate that conventional fisheries management leads to much more extensive and pervasive overfishing than currently realized; harvests are, on average, over twenty-fold too large to be fully sustainable. In general, our approach involves not only the sustainability of fisheries and related resources but also the sustainability of the ecosystems and evosystems in which they occur. Using macroecological patterns accomplishes four important goals: 1) Macroecology becomes one of the interdisciplinary components of management. 2) Sustainability becomes an option for harvests from populations of individual species, species groups, ecosystems, and the entire marine environment. 3) Policies and goals are reality-based, holistic, or fully systemic; they account for ecological as well as evolutionary factors and dynamics (including management itself). 4) Numerous management questions can be addressed.

Genetic heterogeneity and ploidy level analysis among different gynogenetic clones of the polyploid gibel carp

Background: Some triploid and tetraploid clones have been identified in the gynogenetic gibel carp, Carassius auratus gibelio Bloch, by karyotypic and cytologic analyses over many years. Further, 5-20% males and karyotypic diversity have been found among their natural and artificial populations. However, the DNA contents and the relation to their ploidy level and chromosome numbers have not been ascertained, and whether normal meiosis occurs in spermatogenesis needs to be determined in the different clones. Methods: The sampled blood cells or sperms were mixed with blood cells from chicken or individual gibel carp and fixed in 70% pre-cooled ethanol overnight at 4degreesC. The mixed cell pellets were washed 2-3 times in 1x phosphate buffered saline and then resuspended in the solution containing 0.5% pepsin and 0.1 M HCl. DNA was stained with propidium iodide solution (40 mug/mL) containing 4 kU/ml RNase. The measurements of DNA contents were performed with Phoenix Flow Systems. Results: Triploid clones A, E, F, and P had almost equal DNA content, but triploid clone D had greater DNA content than did the other four triploid clones. DNA content of clone M (7.01 +/- 0.15 pg/nucleus) was almost equal to the DNA content of clone D (5-38 +/- 0.06 pg/nucleus) plus the DNA content of common carp sperm (1.64 +/- 0.02 pg/nucleus). The DNA contents of sperms from clones A, P, and D were half of their blood cells, suggesting that normal meiosis occurs in spermatogenesis. Conclusions: Flow cytometry is a powerful method to analyze genetic heterogeneity and ploidy level among different gynogenetic clones of polyploid gibel carp. Through this study, four questions have been answered. (a) The DNA content correlation among the five triploid clones and one multiple tetraploid clone was revealed in the gibel carp, and the contents increased with not only the ploidy level but also the chromosome number. (b) Mean DNA content was 0.052 pg in six extra chromosomes of clone D, which was higher than that of each chromosome in clones A, E, F, and P (about 0.032 pg/ chromosome). This means that the six extra chromosomes are larger chromosomes. (c) Normal meiosis occurred during spermatogenesis of the gibel carp, because DNA contents of the sperms from clones A, P, and D were almost half of that in their blood cells. (d) Multiple tetraploid clone M (7.01 +/- 0.15 pg/nucleus) contained the complete genome of clone D (5.38 +/- 0.06 pg/nucleus) and the genome of common carp sperm (1.64 +/- 0.02 pg/nucleus). Cytometry Part A 56A:46-52, 2003. (C) 2003 Wiley-Liss, Inc.

Layer-by-layer assembly of gelatin

Electrostatic assembly of one species can be realized using gelatin as a polyampholyte. Under suitable conditions where the electrostatic attraction and repulsion were both significant and in balance, linear growth of multilayers driven by electrostatic interactions was sustained over many successive assembly steps, and the maximum amount of adsorption of each layer was reached when the solution pH was around the isoelectric point. The rearrangement of the adsorbed chains after drying was confirmed by contact angle analysis. In addition with only one species involved, the assembled thin films should be chemically uniform rather than layered.

Articulated Pose Estimation in a Learned Smooth Space of Feasible Solutions

A learning based framework is proposed for estimating human body pose from a single image. Given a differentiable function that maps from pose space to image feature space, the goal is to invert the process: estimate the pose given only image features. The inversion is an ill-posed problem as the inverse mapping is a one to many process. Hence multiple solutions exist, and it is desirable to restrict the solution space to a smaller subset of feasible solutions. For example, not all human body poses are feasible due to anthropometric constraints. Since the space of feasible solutions may not admit a closed form description, the proposed framework seeks to exploit machine learning techniques to learn an approximation that is smoothly parameterized over such a space. One such technique is Gaussian Process Latent Variable Modelling. Scaled conjugate gradient is then used find the best matching pose in the space of feasible solutions when given an input image. The formulation allows easy incorporation of various constraints, e.g. temporal consistency and anthropometric constraints. The performance of the proposed approach is evaluated in the task of upper-body pose estimation from silhouettes and compared with the Specialized Mapping Architecture. The estimation accuracy of the Specialized Mapping Architecture is at least one standard deviation worse than the proposed approach in the experiments with synthetic data. In experiments with real video of humans performing gestures, the proposed approach produces qualitatively better estimation results.

Traffic Characteristics and Communication Patterns in Blogosphere

We present a thorough characterization of the access patterns in blogspace -- a fast-growing constituent of the content available through the Internet -- which comprises a rich interconnected web of blog postings and comments by an increasingly prominent user community that collectively define what has become known as the blogosphere. Our characterization of over 35 million read, write, and administrative requests spanning a 28-day period is done from three different blogosphere perspectives. The server view characterizes the aggregate access patterns of all users to all blogs; the user view characterizes how individual users interact with blogosphere objects (blogs); the object view characterizes how individual blogs are accessed. Our findings support two important conclusions. First, we show that the nature of interactions between users and objects is fundamentally different in blogspace than that observed in traditional web content. Access to objects in blogspace could be conceived as part of an interaction between an author and its readership. As we show in our work, such interactions range from one-to-many "broadcast-type" and many-to-one "registration-type" communication between an author and its readers, to multi-way, iterative "parlor-type" dialogues among members of an interest group. This more-interactive nature of the blogosphere leads to interesting traffic and communication patterns, which are different from those observed in traditional web content. Second, we identify and characterize novel features of the blogosphere workload, and we investigate the similarities and differences between typical web server workloads and blogosphere server workloads. Given the increasing share of blogspace traffic, understanding such differences is important for capacity planning and traffic engineering purposes, for example.

Large-Scale Neural Systems for Vision and Cognition

— Consideration of how people respond to the question What is this? has suggested new problem frontiers for pattern recognition and information fusion, as well as neural systems that embody the cognitive transformation of declarative information into relational knowledge. In contrast to traditional classification methods, which aim to find the single correct label for each exemplar (This is a car), the new approach discovers rules that embody coherent relationships among labels which would otherwise appear contradictory to a learning system (This is a car, that is a vehicle, over there is a sedan). This talk will describe how an individual who experiences exemplars in real time, with each exemplar trained on at most one category label, can autonomously discover a hierarchy of cognitive rules, thereby converting local information into global knowledge. Computational examples are based on the observation that sensors working at different times, locations, and spatial scales, and experts with different goals, languages, and situations, may produce apparently inconsistent image labels, which are reconciled by implicit underlying relationships that the network’s learning process discovers. The ARTMAP information fusion system can, moreover, integrate multiple separate knowledge hierarchies, by fusing independent domains into a unified structure. In the process, the system discovers cross-domain rules, inferring multilevel relationships among groups of output classes, without any supervised labeling of these relationships. In order to self-organize its expert system, the ARTMAP information fusion network features distributed code representations which exploit the model’s intrinsic capacity for one-to-many learning (This is a car and a vehicle and a sedan) as well as many-to-one learning (Each of those vehicles is a car). Fusion system software, testbed datasets, and articles are available from http://cns.bu.edu/techlab.

Measuring fast gene dynamics in single cells with time-lapse luminescence microscopy.

Time-lapse fluorescence microscopy is an important tool for measuring in vivo gene dynamics in single cells. However, fluorescent proteins are limited by slow chromophore maturation times and the cellular autofluorescence or phototoxicity that arises from light excitation. An alternative is luciferase, an enzyme that emits photons and is active upon folding. The photon flux per luciferase is significantly lower than that for fluorescent proteins. Thus time-lapse luminescence microscopy has been successfully used to track gene dynamics only in larger organisms and for slower processes, for which more total photons can be collected in one exposure. Here we tested green, yellow, and red beetle luciferases and optimized substrate conditions for in vivo luminescence. By combining time-lapse luminescence microscopy with a microfluidic device, we tracked the dynamics of cell cycle genes in single yeast with subminute exposure times over many generations. Our method was faster and in cells with much smaller volumes than previous work. Fluorescence of an optimized reporter (Venus) lagged luminescence by 15-20 min, which is consistent with its known rate of chromophore maturation in yeast. Our work demonstrates that luciferases are better than fluorescent proteins at faithfully tracking the underlying gene expression.

Marine ecosystem response to the Atlantic Multidecadal Oscillation

Against the backdrop of warming of the Northern Hemisphere it has recently been acknowledged that North Atlantic temperature changes undergo considerable variability over multidecadal periods. The leading component of natural low-frequency temperature variability has been termed the Atlantic Multidecadal Oscillation (AMO). Presently, correlative studies on the biological impact of the AMO on marine ecosystems over the duration of a whole AMO cycle (~60 years) is largely unknown due to the rarity of continuously sustained biological observations at the same time period. To test whether there is multidecadal cyclic behaviour in biological time-series in the North Atlantic we used one of the world's longest continuously sustained marine biological time-series in oceanic waters, long-term fisheries data and historical records over the last century and beyond. Our findings suggest that the AMO is far from a trivial presence against the backdrop of continued temperature warming in the North Atlantic and accounts for the second most important macro-trend in North Atlantic plankton records; responsible for habitat switching (abrupt ecosystem/regime shifts) over multidecadal scales and influences the fortunes of various fisheries over many centuries.

A general framework for aquatic biogeochemical models

One of the most of challenging steps in the development of coupled hydrodynamic-biogeochemical models is the combination of multiple, often incompatible computer codes that describe individual physical, chemical, biological and geological processes. This “coupling” is time-consuming, error-prone, and demanding in terms of scientific and programming expertise. The open source, Fortran-based Framework for Aquatic Biogeochemical Models addresses these problems by providing a consistent set of programming interfaces through which hydrodynamic and biogeochemical models communicate. Models are coded once to connect to FABM, after which arbitrary combinations of hydrodynamic and biogeochemical models can be made. Thus, a biogeochemical model code works unmodified within models of a chemostat, a vertically structured water column, and a three-dimensional basin. Moreover, complex biogeochemistry can be distributed over many compact, self-contained modules, coupled at run-time. By enabling distributed development and user-controlled coupling of biogeochemical models, FABM enables optimal use of the expertise of scientists, programmers and end-users.