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Ice rises or ice rumples act as ice-shelf pinning points that can play an important role in regulating the ice discharge of marine outlet glaciers. As an example, the observed recent gradual ungrounding of the ice shelf of West Antarctica's Thwaites Glacier from its last pinning points likely diminished the buttressing effect of the ice shelf and thus contributed to the destabilization of the outlet. Here we use an idealized experimental setting to simulate the response of a marine outlet glacier resting on a landward down-sloping (retrograde) bed to a stepwise ungrounding of its ice shelf from a topographic high and a subsequent re-grounding. We show that the glacier retreat down the retrograde bed, induced by the loss in pinning-point buttressing, can be unstable and irreversible given a relatively deep subglacial bed depression. In this case, glacier retreat and re-advance show a hysteretic behavior, and if the bed depression is sufficiently deep, the glacier does not recover but remains locked in its retreated state. Conversely, reversibility requires a sufficiently shallow bed depression. Based on a simple flux balance analysis, we argue that the combination of a deep bed depression and limited ice-shelf buttressing hampers grounding-line re-advance due to the dominant and highly non-linear influence of the bed depth on the ice discharge across the grounding line. We conclude that outlets that rest on a deep bed depression and are weakly buttressed, such as Thwaites Glacier, are more susceptible to abrupt and irreversible retreat than more strongly buttressed glaciers on more moderate retrograde slopes, such as Pine Island Glacier. Our findings further suggest that the (ir)reversibility of large-scale grounding-line retreat may be strongly affected by calving-front migration and associated changes in ice-shelf buttressing.

Fusarium oxysporum f. sp. fragariae (Fof) race 1 is avirulent on cultivars with the dominant resistance gene FW1, while Fof race 2 is virulent on FW1-resistant cultivars. We hypothesized there was a gene-for-gene interaction between a gene at the FW1 locus and an avirulence gene (AvrFW1) in Fof race 1. To identify a candidate AvrFW1, we compared genomes of 24 Fof race 1 and three Fof race 2 isolates. We found one candidate gene that was present in race 1, was absent in race 2, was highly expressed in planta, and was homologous to a known effector, secreted in xylem 6 (SIX6). We knocked out SIX6 in two Fof race 1 isolates by homologous recombination. All SIX6 knockout transformants (ΔSIX6) gained virulence on FW1/fw1 cultivars, whereas ectopic transformants and the wildtype isolates remained avirulent. ΔSIX6 isolates were quantitatively less virulent on FW1/fw1 cultivars Fronteras and San Andreas than fw1/fw1 cultivars. Seedlings from an FW1/fw1 × fw1/fw1 population were genotyped for FW1 and tested for susceptibility to a SIX6 knockout isolate. Results suggested that additional minor-effect quantitative resistance genes could be present at the FW1 locus. This work demonstrates that SIX6 acts as an avirulence factor interacting with a resistance gene at the FW1 locus. The identification of AvrFW1 enables surveillance for Fof race 2 and provides insight into the mechanisms of FW1-mediated resistance. [Graphic: see text] Copyright © 2024 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license.

Abstract The annual production of strawberry has increased by one million tonnes in the US and 8.4 million tonnes worldwide since 1960. Here we show that the US expansion was driven by genetic gains from Green Revolution breeding and production advances that increased yields by 2,755%. Using a California population with a century-long breeding history and phenotypes of hybrids observed in coastal California environments, we estimate that breeding has increased fruit yields by 2,974-6,636%, counts by 1,454-3,940%, weights by 228-504%, and firmness by 239-769%. Using genomic prediction approaches, we pinpoint the origin of the Green Revolution to the early 1950s and uncover significant increases in additive genetic variation caused by transgressive segregation and phenotypic diversification. Lastly, we show that the most consequential Green Revolution breeding breakthrough was the introduction of photoperiod-insensitive, PERPETUAL FLOWERING hybrids in the 1970s that doubled yields and drove the dramatic expansion of strawberry production in California.

Abstract Verticillium wilt (VW), a devastating vascular wilt disease of strawberry (Fragaria × ananassa), has caused economic losses for nearly a century. This disease is caused by the soil‐borne pathogen Verticillium dahliae, which occurs nearly worldwide and causes disease in numerous agriculturally important plants. The development of VW‐resistant cultivars is critically important for the sustainability of strawberry production. We previously showed that a preponderance of the genetic resources (asexually propagated hybrid individuals) preserved in public germplasm collections were moderately to highly susceptible and that genetic gains for increased resistance to VW have been negligible over the last 60 years. To more fully understand the challenges associated with breeding for increased quantitative resistance to this pathogen, we developed and phenotyped a training population of hybrids (n=564) among elite parents with a wide range of resistance phenotypes. When these data were combined with training data from a population of elite and exotic hybrids (n=386), genomic prediction accuracies of 0.47–0.48 were achieved and were predicted to explain 70%–75% of the additive genetic variance for resistance. We concluded that breeding values for resistance to VW can be predicted with sufficient accuracy for effective genomic selection with routine updating of training populations.

Abstract Genomic prediction in breeding populations containing hundreds to thousands of parents and seedlings is prohibitively expensive with current high‐density genetic marker platforms designed for strawberry. We developed mid‐density panels of molecular inversion probes (MIPs) to be deployed with the “DArTag” marker platform to provide a low‐cost, high‐throughput genotyping solution for strawberry genomic prediction. In total, 7742 target single nucleotide polymorphism (SNP) regions were used to generate MIP assays that were tested with a screening panel of 376 octoploid Fragaria accessions. We evaluated the performance of DArTag assays based on genotype segregation, amplicon coverage, and their ability to produce subgenome‐specific amplicon alignments to the FaRR1 assembly and subsequent alignment‐based variant calls with strong concordance to DArT's alignment‐free, count‐based genotype reports. We used a combination of marker performance metrics and physical distribution in the FaRR1 assembly to select 3K and 5K production panels for genotyping of large strawberry populations. We show that the 3K and 5K DArTag panels are able to target and amplify homologous alleles within subgenomic sequences with low‐amplification bias between reference and alternate alleles, supporting accurate genotype calling while producing marker genotypes that can be treated as functionally diploid for quantitative genetic analysis. The 3K and 5K target SNPs show high levels of polymorphism in diverse F. × ananassa germplasm and UC Davis cultivars, with mean pairwise diversity (π) estimates of 0.40 and 0.32 and mean heterozygous genotype frequencies of 0.35 and 0.33, respectively.

The timescales of the flow and retreat of Greenland's and Antarctica's outlet glaciers and their potential instabilities are arguably the largest uncertainty in future sea-level projections. Here we derive a scaling relation that allows the comparison of the timescales of observed complex ice flow fields with geometric similarity. The scaling relation is derived under the assumption of fast, laterally confined, geometrically similar outlet-glacier flow over a slippery bed, i.e., with negligible basal friction. According to the relation, the time scaling of the outlet flow is determined by the product of the inverse of (1) the fourth power of the width-to-length ratio of its confinement, (2) the third power of the confinement depth and (3) the temperature-dependent ice softness. For the outflow at the grounding line of streams with negligible basal friction, this means that the volume flux is proportional to the ice softness and the bed depth, but goes with the fourth power of the gradient of the bed and with the fifth power of the width of the stream. We show that the theoretically derived scaling relation is supported by the observed velocity scaling of outlet glaciers across Greenland as well as by idealized numerical simulations of marine ice-sheet instabilities (MISIs) as found in Antarctica. Assuming that changes in the ice-flow velocity due to ice-dynamic imbalance are proportional to the equilibrium velocity, we combine the scaling relation with a statistical analysis of the topography of 13 MISI-prone Antarctic outlets. Under these assumptions, the timescales in response to a potential destabilization are fastest for Thwaites Glacier in West Antarctica and Mellor, Ninnis and Cook Glaciers in East Antarctica; between 16 and 67 times faster than for Pine Island Glacier. While the applicability of our results is limited by several strong assumptions, the utilization and potential further development of the presented scaling approach may help to constrain timescale estimates of outlet-glacier flow, augmenting the commonly exploited and comparatively computationally expensive approach of numerical modeling.

Crop production is becoming an increasing challenge as the global population grows and the climate changes. Modern cultivated crop species are selected for productivity under optimal growth environments and have often lost genetic variants that could allow them to adapt to diverse, and now rapidly changing, environments. These genetic variants are often present in their closest wild relatives, but so are less desirable traits. How to preserve and effectively utilize the rich genetic resources that crop wild relatives offer while avoiding detrimental variants and maladaptive genetic contributions is a central challenge for ongoing crop improvement. This Essay explores this challenge and potential paths that could lead to a solution.

Owing to global warming and particularly high regional ocean warming, both Thwaites and Pine Island Glaciers in the Amundsen region of the Antarctic Ice Sheet could lose their buttressing ice shelves over time. We analyse the possible consequences using the parallel ice sheet model (PISM), applying a simple cliff-calving parameterization and an ice mélange-buttressing model. We find that the instantaneous loss of ice-shelf buttressing, due to enforced ice-shelf melting, initiates grounding-line retreat and triggers marine ice sheet instability (MISI). As a consequence, the grounding line progresses into the interior of the West Antarctic Ice Sheet and leads to a sea level contribution of 0.6 m within 100 a. By subjecting the exposed ice cliffs to cliff calving using our simplified parameterization, we also analyse marine ice cliff instability (MICI). In our simulations it can double or even triple the sea level contribution depending on the only loosely constrained parameter that determines the maximum cliff-calving rate. The speed of MICI depends on this upper bound of the calving rate, which is given by the ice mélange buttressing the glacier. However, stabilization of MICI may occur for geometric reasons. Because the embayment geometry changes as MICI advances into the interior of the ice sheet, the upper bound on calving rates is reduced and the progress of MICI is slowed down. Although we cannot claim that our simulations bear relevant quantitative estimates of the effect of ice-mélange buttressing on MICI, the mechanism has the potential to stop the instability. Further research is needed to evaluate its role for the past and future evolution of the Antarctic Ice Sheet.

Basal ice-shelf melting is the key driver of Antarctica's increasing sea-level contribution. In diminishing the buttressing force of the ice shelves that fringe the ice sheet, the melting increases the ice discharge into the ocean. Here we contrast the influence of basal melting in two different ice-shelf regions on the time-dependent response of an isothermal, inherently buttressed ice-sheet-shelf system. In the idealized numerical simulations, the basal-melt perturbations are applied close to the grounding line in the ice-shelf's (1) ice-stream region, where the ice shelf is fed by the fastest ice masses that stream through the upstream bed trough and (2) shear margins, where the ice flow is slower. The results show that melting below one or both of the shear margins can cause a decadal to centennial increase in ice discharge that is more than twice as large compared to a similar perturbation in the ice-stream region. We attribute this to the fact that melt-induced ice-shelf thinning in the central grounding-line region is attenuated very effectively by the fast flow of the central ice stream. In contrast, the much slower ice dynamics in the lateral shear margins of the ice shelf facilitate sustained ice-shelf thinning and thereby foster buttressing reduction. Regardless of the melt location, a higher melt concentration toward the grounding line generally goes along with a stronger response. Our results highlight the vulnerability of outlet glaciers to basal melting in stagnant, buttressing-relevant ice-shelf regions, a mechanism that may gain importance under future global warming.

Abstract The development of strawberry (Fragaria × ananassa Duchesne ex Rozier) cultivars resistant to Phytophthora crown rot (PhCR), a devastating disease caused by the soil‐borne pathogen Phytophthora cactorum (Lebert & Cohn) J. Schröt., has been challenging partly because the resistance phenotypes are quantitative and only moderately heritable. To develop deeper insights into the genetics of resistance and build the foundation for applying genomic selection, a genetically diverse training population was screened for resistance to California isolates of the pathogen. Here we show that genetic gains in breeding for resistance to PhCR have been negligible (3% of the cultivars tested were highly resistant and none surpassed early 20th century cultivars). Narrow‐sense genomic heritability for PhCR resistance ranged from 0.41 to 0.75 among training population individuals. Using multivariate genome‐wide association studies (GWAS), we identified a large‐effect locus (predicted to be RPc2) that explained 43.6–51.6% of the genetic variance, was necessary but not sufficient for resistance, and was associated with calcium channel and other candidate genes with known plant defense functions. The addition of underutilized gene bank resources to our training population doubled additive genetic variance, increased the accuracy of genomic selection, and enabled the discovery of individuals carrying favorable alleles that are either rare or not present in modern cultivars. The incorporation of an RPc2‐associated single‐nucleotide polymorphism (SNP) as a fixed effect increased genomic prediction accuracy from 0.40 to 0.55. Finally, we show that parent selection using genomic‐estimated breeding values, genetic variances, and cross usefulness holds promise for enhancing resistance to PhCR in strawberry.

AbstractMany important traits in plants, animals, and microbes are polygenic and challenging to improve through traditional marker-assisted selection. Genomic prediction addresses this by incorporating all genetic data in a mixed model framework. The primary method for predicting breeding values is genomic best linear unbiased prediction, which uses the realized genomic relationship or kinship matrix (K σ g 2 h g 2K σ g 2 h g 2ZKH K ASV σ g 2 h g 2

We present the result of the third Marine Ice Sheet Model Intercomparison Project, MISMIP+. MISMIP+ is intended to be a benchmark for ice-flow models which include fast sliding marine ice streams and floating ice shelves and in particular a treatment of viscous stress that is sufficient to model buttressing, where upstream ice flow is restrained by a downstream ice shelf. A set of idealized experiments first tests that models are able to maintain a steady state with the grounding line located on a retrograde slope due to buttressing and then explore scenarios where a reduction in that buttressing causes ice stream acceleration, thinning, and grounding line retreat. The majority of participating models passed the first test and then produced similar responses to the loss of buttressing. We find that the most important distinction between models in this particular type of simulation is in the treatment of sliding at the bed, with other distinctions – notably the difference between the simpler and more complete treatments of englacial stress but also the differences between numerical methods – taking a secondary role.

Abstract Reliable phenotyping methods that are simple to operate and inexpensive to deploy are critical for studying quantitative traits in plants. Traditional fruit shape phenotyping relies on human raters or 2D analyses to assess form, e.g., size and shape. Systems for 3D imaging using multi‐view stereo have been implemented, but frequently rely on commercial software and/or specialized hardware, which can lead to limitations in accessibility and scalability. We present a complete system constructed of consumer‐grade components for capturing, calibrating, and reconstructing the 3D form of small‐to‐moderate sized fruits and tubers. Data acquisition and image capture sessions are 9 seconds to capture 60 images. The initial prototype cost was $1600 USD. We measured accuracy by comparing reconstructed models of 3D printed ground truth objects to the original digital files of those same ground truth objects. The R2 between length of the primary, secondary, and tertiary axes, volume, and surface area of the ground‐truth object and the reconstructed models was >0.97 and root‐mean square error (RMSE) was < 3 mm for objects without locally concave regions. Measurements from 1 mm and 2 mm resolution reconstructions were consistent (R2 > 0.99). Qualitative assessments were performed on 48 fruit and tubers, including 18 strawberries, 12 potatoes, five grapes, seven peppers, and four Bosc and two red Anjou pears. Our proposed phenotyping system is fast, relatively low cost, and has demonstrated accuracy for certain shape classes, and could be used for the 3D analysis of fruit form.

AbstractGray mold, a disease of strawberry (FragariaananassaBotrytis cinerea

The development of genome-informed methods for identifying quantitative trait loci (QTL) and studying the genetic basis of quantitative variation in natural and experimental populations has been driven by advances in high-throughput genotyping. For many complex traits, the underlying genetic variation is caused by the segregation of one or more 'large-effect' loci, in addition to an unknown number of loci with effects below the threshold of statistical detection. The large-effect loci segregating in populations are often necessary but not sufficient for predicting quantitative phenotypes. They are, nevertheless, important enough to warrant deeper study and direct modelling in genomic prediction problems. We explored the accuracy of statistical methods for estimating the fraction of marker-associated genetic variance (p) and heritability ([Formula: see text]) for large-effect loci underlying complex phenotypes. We found that commonly used statistical methods overestimate p and [Formula: see text]. The source of the upward bias was traced to inequalities between the expected values of variance components in the numerators and denominators of these parameters. Algebraic solutions for bias-correcting estimates of p and [Formula: see text] were found that only depend on the degrees of freedom and are constant for a given study design. We discovered that average semivariance methods, which have heretofore not been used in complex trait analyses, yielded unbiased estimates of p and [Formula: see text], in addition to best linear unbiased predictors of the additive and dominance effects of the underlying loci. The cryptic bias problem described here is unrelated to selection bias, although both cause the overestimation of p and [Formula: see text]. The solutions we described are predicted to more accurately describe the contributions of large-effect loci to the genetic variation underlying complex traits of medical, biological, and agricultural importance.

Recent observations and ice-dynamic modeling suggest that a marine ice-sheet instability (MISI) might have been triggered in West Antarctica. The corresponding outlet glaciers, Pine Island Glacier (PIG) and Thwaites Glacier (TG), showed significant retreat during at least the last 2 decades. While other regions in Antarctica have the topographic predisposition for the same kind of instability, it is so far unclear how fast these instabilities would unfold if they were initiated. Here we employ the concept of similitude to estimate the characteristic timescales of several potentially MISI-prone outlet glaciers around the Antarctic coast. Our results suggest that TG and PIG have the fastest response time of all investigated outlets, with TG responding about 1.25 to 2 times as fast as PIG, while other outlets around Antarctica would be up to 10 times slower if destabilized. These results have to be viewed in light of the strong assumptions made in their derivation. These include the absence of ice-shelf buttressing, the one-dimensionality of the approach and the uncertainty of the available data. We argue however that the current topographic situation and the physical conditions of the MISI-prone outlet glaciers carry the information of their respective timescale and that this information can be partially extracted through a similitude analysis.

Cultivated strawberry has a compelling domestication history: the earliest cultivars originated in western Europe as spontaneous hybrids between non-sympatric wild species imported from the New World. The origin of those early hybrids went undiscovered for nearly a half century before conscious domestication began. Here, Pincot, Ledda, Feldmann et al

Iron oxide grain coatings in red sandstones contain trace metals that are released upon dissolution of the coatings. Analyses by ICP-MS following acid leaching of the grain coatings show that the dissolved metals can constitute an ore-forming fluid, as hypothesized in models for sandstone-hosted ore deposits. Median compositions of 37 samples, mostly of Triassic to Devonian age, from across Britain and Ireland are 6.3 ppm copper, 2.4 ppm cobalt, 10.1 ppm vanadium, and 0.3 ppm uranium. These contents at the basin scale are adequate to form the observed range of ore deposits in red beds. The migration of hydrocarbons or brines can cause the dissolution of grain coatings and contributes to controlling the distribution of ore deposits. Future measurements should test red beds derived from uplifted, mineralized plate margins, in which sandstones may be preloaded with ore metals.

Abstract Understanding the genetic basis of plant traits requires comprehensive and quantitative descriptions of the phenotypic variation that exists within populations. Cameras and other sensors have made high‐throughput phenotyping possible, but image‐based phenotyping procedures involve a step where a researcher selects the traits to be measured. This feature selection step is inherently prone to human biases. Recently, a set of phenotyping approaches, which are referred to collectively as latent phenotyping techniques, have arisen in the literature. Latent phenotyping techniques isolate a latent source of variance in the data, such as stress or genotype, and then quantify the effect of this latent source of variance using latent variables without defining any conventional traits. In this review, we discuss the differences between, and challenges of, both traditional and latent phenotyping.

Allo-octoploid cultivated strawberry (Fragaria × ananassa) originated through a combination of polyploid and homoploid hybridization, domestication of an interspecific hybrid lineage, and continued admixture of wild species over the last 300 years. While genes appear to flow freely between the octoploid progenitors, the genome structures and diversity of the octoploid species remain poorly understood. The complexity and absence of an octoploid genome frustrated early efforts to study chromosome evolution, resolve subgenomic structure, and develop a single coherent linkage group nomenclature. Here, we show that octoploid Fragaria species harbor millions of subgenome-specific DNA variants. Their diversity was sufficient to distinguish duplicated (homoeologous and paralogous) DNA sequences and develop 50K and 850K SNP genotyping arrays populated with co-dominant, disomic SNP markers distributed throughout the octoploid genome. Whole-genome shotgun genotyping of an interspecific segregating population yielded 1.9M genetically mapped subgenome variants in 5,521 haploblocks spanning 3,394 cM in F. chiloensis subsp. lucida, and 1.6M genetically mapped subgenome variants in 3,179 haploblocks spanning 2,017 cM in F. × ananassa. These studies provide a dense genomic framework of subgenome-specific DNA markers for seamlessly cross-referencing genetic and physical mapping information and unifying existing chromosome nomenclatures. Using comparative genomics, we show that geographically diverse wild octoploids are effectively diploidized, nearly completely collinear, and retain strong macro-synteny with diploid progenitor species. The preservation of genome structure among allo-octoploid taxa is a critical factor in the unique history of garden strawberry, where unimpeded gene flow supported its origin and domestication through repeated cycles of interspecific hybridization.

Computing approaches in the optical domain would allow for ultra-fast signaling and ultra-high bandwidth capabilities. Here, Feldmann et al. demonstrate a photonic abacus, which provides multistate compute-and store operation by integrating phase-change materials with nanophotonic chips.

>Here we report on a cyclic, physical ice-discharge instability in the Parallel Ice Sheet Model, simulating the flow of a three-dimensional, inherently buttressed ice-sheet-shelf system which periodically surges on a millennial timescale. The thermomechanically coupled model on 1 km horizontal resolution includes an enthalpy-based formulation of the thermodynamics, a nonlinear stress-balance-based sliding law and a very simple subglacial hydrology. The simulated unforced surging is characterized by rapid ice streaming through a bed trough, resulting in abrupt discharge of ice across the grounding line which is eventually calved into the ocean. We visualize the central feedbacks that dominate the subsequent phases of ice buildup, surge and stabilization which emerge from the interaction between ice dynamics, thermodynamics and the subglacial till layer. Results from the variation of surface mass balance and basal roughness suggest that ice sheets of medium thickness may be more susceptible to surging than relatively thin or thick ones for which the surge feedback loop is damped. We also investigate the influence of different basal sliding laws (ranging from purely plastic to nonlinear to linear) on possible surging. The presented mechanisms underlying our simulations of self-maintained, periodic ice growth and destabilization may play a role in large-scale ice-sheet surging, such as the surging of the Laurentide Ice Sheet, which is associated with Heinrich events, and ice-stream shutdown and reactivation, such as observed in the Siple Coast region of West Antarctica.

The concept of similitude is commonly employed in the fields of fluid dynamics and engineering but rarely used in cryospheric research. Here we apply this method to the problem of ice flow to examine the dynamic similitude of isothermal ice sheets in shallow-shelf approximation against the scaling of their geometry and physical parameters. Carrying out a dimensional analysis of the stress balance we obtain dimensionless numbers that characterize the flow. Requiring that these numbers remain the same under scaling we obtain conditions that relate the geometric scaling factors, the parameters for the ice softness, surface mass balance and basal friction as well as the ice-sheet intrinsic response time to each other. We demonstrate that these scaling laws are the same for both the (two-dimensional) flow-line case and the three-dimensional case. The theoretically predicted ice-sheet scaling behavior agrees with results from numerical simulations that we conduct in flow-line and three-dimensional conceptual setups. We further investigate analytically the implications of geometric scaling of ice sheets for their response time. With this study we provide a framework which, under several assumptions, allows for a fundamental comparison of the ice-dynamic behavior across different scales. It proves to be useful in the design of conceptual numerical model setups and could also be helpful for designing laboratory glacier experiments. The concept might also be applied to real-world systems, e.g., to examine the response times of glaciers, ice streams or ice sheets to climatic perturbations.

The initiation of a marine ice-sheet instability (MISI) is generally discussed from the ocean side of the ice sheet. It has been shown that the reduction in ice-shelf buttressing and softening of the coastal ice can destabilize a marine ice sheet if the bedrock is sloping upward towards the ocean. Using a conceptional flow-line geometry, we investigate the possibility of whether a MISI can be triggered from the direction of the ice divide as opposed to coastal forcing and explore the interaction between connected basins. We find that the initiation of a MISI in one basin can induce a destabilization in the other. The underlying mechanism of basin interaction is based on dynamic thinning and a consecutive motion of the ice divide which induces a thinning in the adjacent basin and a successive initiation of the instability. Our simplified and symmetric topographic setup allows scaling both the geometry and the transition time between both instabilities. We find that the ice profile follows a universal shape that is scaled with the horizontal extent of the ice sheet and that the same exponent of 1/2 applies for the scaling relation between central surface elevation and horizontal extent as in the pure shallow ice approximation (Vialov profile). Altering the central bed elevation, we find that the extent of grounding-line retreat in one basin determines the degree of interaction with the other. Different scenarios of basin interaction are discussed based on our modeling results as well as on a conceptual flux-balance analysis. We conclude that for the three-dimensional case, the possibility of drainage basin interaction on timescales on the order of 1 kyr or larger cannot be excluded and hence needs further investigation.