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Ryan, M.

Publications and source records attributed to Ryan, M..

At least 19 recordsLinked to original sources

Resolving platypus behaviour from accelerometry: frequency-domain features improve detection of rhythmic behaviours in hydrodynamically challenging aquatic environments

Developing robust methods to quantify how animals allocate time across behaviours is essential for understanding energy use, habitat requirements, and responses to environmental change. For cryptic, semi-aquatic mammals such as the platypus, direct observation is difficult, creating a reliance on remote biologging approaches that can reliably infer behaviour in the wild. However, aquatic environments can both smooth acceleration signals through hydrodynamic damping and introduce noise from water movement, turbulence, and drag, potentially obscuring behavioural differences of similar magnitudes. We tested whether progressively incorporating biomechanical and frequency-domain (FFT-derived) predictors improved behavioural classification in hydrodynamically challenging aquatic environments. Tri-axial accelerometers were deployed on four ex situ platypuses, with synchronised video observations used to validate behaviour. From the acceleration data, we derived three predictor classes of increasing complexity: summary statistics describing activity level, engineered biomechanical variables capturing posture and body orientation, and FFT-derived features describing movement rhythm. These predictors were progressively incorporated into Random Forest models to classify five behaviours: burrow resting, surface resting, grooming, travelling/foraging, and diving. Model performance improved with increasing predictor complexity, although gains were behaviour specific. FFT-derived features substantially improved classification of rhythmic behaviours such as diving and foraging, while engineered biomechanical predictors improved grooming detection. In contrast, resting behaviours, particularly surface resting, showed little improvement. Overall accuracy increased from [~]75% to [~]88% when frequency-domain features were included. Misclassification was greatest among behaviours with overlapping or low-amplitude signals, and cross-individual validation revealed reduced model generalisability, indicating that individual variation in movement patterns constrained transferability. Incorporating frequency-domain features substantially improved behavioural classification in platypuses, particularly for rhythmic behaviours such as diving and foraging. This study provides the first validated accelerometry-based behavioural classification framework for the species and highlights the importance of matching predictor selection to behavioural mechanics. More broadly, the approach offers a transferable framework for aquatic and semi-aquatic taxa.

animal behavior and cognition↗

Splicing of ultraconserved poison exons controls mitotic fidelity and stem cell viability

SR proteins are essential splicing regulators whose expression is controlled in part through poison exons (PEs) -- ultraconserved non-coding exons that trigger nonsense-mediated decay -- yet the biological functions of these elements remain undefined. Here, we show that homozygous deletion of SRSF3-PE or TRA2{beta}-PE is selected against in mouse embryos and human induced pluripotent stem cells (iPSCs), and that conditional PE deletion causes apoptotic death in iPSCs but is tolerated in post-mitotic neurons, revealing a proliferative-state-specific requirement. Mechanistically, PE deletion elevates SR protein levels, triggers widespread splicing dysregulation, and disrupts the correct splicing of a mitotic gene network associated with spindle defects and mitotic errors. These findings establish ultraconserved poison exons as essential regulators of mitotic splicing fidelity and stem cell viability.

molecular biology↗

Genetic factors driving multi-host infection in a core member of the root mycobiota

Core members of the fungal root microbiota include pathogens capable of colonizing multiple hosts, yet the underlying genetic determinants remain unknown. We report that Plectosphaerella cucumerina is a core member of the Arabidopsis thaliana root microbiota displaying high pathogenic potential and multi-host colonization capabilities. Establishment of a Plectosphaerella reference culture collection, followed by whole-genome sequencing of 72 strains reveals subtle phenotypic and genotypic variation that associate with fungal phylogeny, but not host plant identity. Transcriptome profiling of a model P. cucumerina isolate in roots of multiple hosts identifies core and host-specific fungal processes linked to carbon catabolism and root cell wall deconstruction of the hosts. A fungal gene encoding a candidate {beta}-1,3-glucanase (GH64) was identified as a key genetic factor driving infection and disease in plants that diverged 110 million years ago. The gene is enriched in plant-colonizing fungi and consistently functions as a disease determinant in the root pathogen Colletotrichum incanum. We conclude that diverse and tunable fungal repertoires of carbohydrate-active enzymes act as disease determinants and drive multi-host compatibility belowground.

microbiology↗

The potential to breed for genetic legacy effects in sustainable farming systems

Legume crops provide protein-rich food, critical disease breaks in cereal rotations, and contribute to soil fertility through symbiotic nitrogen fixation. However, crop improvement programs typically focus on within-crop performance rather than system-level benefits. We hypothesise that legacy effects (the influence of one crops genotype on subsequent crop performance) are under genetic control and could be leveraged in breeding programs. To test this, we evaluated how 309 genetically diverse mungbean genotypes influence subsequent wheat performance. The mungbean panel was grown, followed by a single wheat cultivar sown in the same plot locations. Remarkably, wheat yield varied by nearly 1 t ha{square}{superscript 1} (2.52-3.49 t ha{square}{superscript 1}) depending solely on the preceding mungbean genotype, with legacy traits displaying moderate heritability (H{superscript 2}: 0.43-0.65), demonstrating untapped genetic potential for breeding. Analyses of mungbean traits, soil properties, and volatile organic compounds implicated root architecture, symbiotic nitrogen fixation and soil microbiome as potential biological drivers of legacy effects. Haplotype mapping identified genomic regions in mungbean associated with wheat yield and protein, revealing trade-offs between within-crop and legacy performance. Genetic simulations using empirically derived marker effects compared genomic selection strategies targeting mungbean yield, wheat yield, or both simultaneously. Balanced selection (50:50 weighting) achieved simultaneous gains in both crops (19.5% and 7.6%), highlighting the opportunity to breed for system-level productivity with reduced input requirements.

plant biology↗

Vocal Interactions Between Singing Humpback Whales (Megaptera novaeangliae)

Humpback whales construct predictably patterned sequences within multi-hour sessions of sound production. These sequences (called "songs") are some of the most structurally and acoustically complex vocal patterns produced by any mammal. Humpback whale song production is highly dynamic in that individuals constantly modify the properties of songs across multiple time scales throughout their adult lives. Past analyses of co-vocalizing humpback whales suggest that singers within earshot of one another may dynamically and interactively adjust their vocalizations in reaction to what they hear. The current study investigated whether singing humpback whales coordinate their production of overlapping sound patterns either by modulating how long they produce similar patterns or by dynamically adjusting the spectral properties of individual sounds and/or sound sequences. Our results show that the vocal adjustments singing humpback whales make when singing in dyads are diverse and appear to depend on the acoustic context. These findings confirm that individual singers flexibly adjust song features in real-time. When pairs of singers adjust song content, they do so in ways that may increase or decrease the acoustic similarity of concurrent sound patterns. No evidence was found of singers adjusting the number or timing of sound pattern repetitions to alter pattern overlap. However, most singers modified how they produced units after a second singer began singing. The current results suggest that co-singing humpback whales attend to the vocal actions of other audible singers and may sometimes modulate song content to initiate social interactions from long distances. Understanding how singing whales vocally interact can clarify the relevance of singing in coordinating vertebrate behavior.

animal behavior and cognition↗

Dried Blood Spot Lipidomics in Diabetes

Comprehensive lipidomic profiling in diabetes has identified disease-associated lipid signatures that may support more personalised monitoring beyond routine glycaemic control. Dried blood spot (DBS) microsampling offers a minimally invasive method for collecting and storing small blood volumes (<50 {micro}L), enabling decentralised and longitudinal sampling. However, its application in lipidomics for cardiometabolic phenotyping remains underexplored. In this study, 34 participants (17 diabetics, 17 non-diabetic controls) self-collected DBS samples using advanced microsampling devices (10 {micro}L, Capitainer(R)B). Lipid extracts were analysed using a validated liquid chromatography-trapped ion mobility-time-of-flight mass spectrometry method optimised for semi-quantitative DBS lipidomics. Multivariate analysis revealed significant differences in lipidomic profiles (n = 432 lipids) between diabetic and non-diabetic individuals (R{superscript 2}Y = 0.82, Q{superscript 2}Y = 0.50, p = 0.05). Key discriminatory lipids (all VIP > 1.44), including HexCer(18:1;O2/24:0), HexCer(18:1;O2/22:0), LPC(16:0), PC(O-34:3), TG(18:0_18:1_20:3), and TG(18:1_18:1_22:6), were consistent with matched venous and capillary plasma samples. Lipid class and structural differences, including elevated long-chain triacylglycerols and reduced lysophosphatidylcholines reflected known dyslipidaemia in diabetes and demonstrate the capacity of DBS to capture biologically meaningful lipid perturbations. Participant-reported outcomes from Problem Areas In Diabetes (PAID-20), Patient Activation Measure (PAM-13), and microsampling perception questionnaires highlighted psychosocial and behavioural insights, and indicated strong support for DBS sampling over venepuncture, citing ease of use and reduced burden. These findings establish DBS lipidomics as a feasible and informative approach for stratifying cardiometabolic disease, supporting broader implementation in remote, personalised monitoring frameworks (Figure 1). O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=77 SRC="FIGDIR/small/656742v1_fig1.gif" ALT="Figure 1"> View larger version (13K): org.highwire.dtl.DTLVardef@c53b4org.highwire.dtl.DTLVardef@14b507forg.highwire.dtl.DTLVardef@191e563org.highwire.dtl.DTLVardef@16af172_HPS_FORMAT_FIGEXP M_FIG O_FLOATNOFigure 1.C_FLOATNO Graphical abstract - Chapter 5. C_FIG

biochemistry↗

Targeting Iron - Respiratory Reciprocity Promotes Bacterial Death

Discovering new bacterial signaling pathways offers unique antibiotic strategies. Here, through an unbiased resistance screen of 3,884 gene knockout strains, we uncovered a previously unknown non-lytic bactericidal mechanism that sequentially couples three transporters and downstream transcription to lethally suppress respiration of the highly virulent P. aeruginosa strain PA14 - one of three species on the WHOs Priority 1: Critical list. By targeting outer membrane YaiW, cationic lacritin peptide N-104 translocates into the periplasm where it ligates outer loops 4 and 2 of the inner membrane transporters FeoB and PotH, respectively, to suppress both ferrous iron and polyamine uptake. This broadly shuts down transcription of many biofilm-associated genes, including ferrous iron-dependent TauD and ExbB1. The mechanism is innate to the surface of the eye and is enhanced by synergistic coupling with thrombin peptide GKY20. This is the first example of an inhibitor of multiple bacterial transporters.

microbiology↗

Large structural variants in KOLF2.1J are unlikely to compromise neurological disease modelling

Gracia-Diaz and colleagues analysed high-density DNA microarray and whole genome sequencing (WGS) data from the KOLF2.1J reference human induced pluripotent stem cell (hiPSC) line1, and report the presence of five high-confidence heterozygous copy number variants (CNVs) at least 100kbp in length2. Since three of these CNVs span coding genes, some of which have been associated with neurodevelopmental disease, the authors raise the concern that these CNVs may compromise the utility of KOLF2.1J for neurological disease modelling. We appreciate their thorough analysis and thoughtful interpretation, and agree that potential users of this line should be made aware of all cases where KOLF2.1J differs from the reference genome. However, we believe that the benefits from the widespread use of KOLF2.1J outweigh the potential risks that might arise from the identified CNVs.

cell biology↗

Postsynaptic mitochondria are positioned to support functional diversity of dendritic spines

Postsynaptic mitochondria are critical to the development, plasticity, and maintenance of synaptic inputs. However, their relationship to synaptic structure and functional activity is unknown. We examined a correlative dataset from ferret visual cortex with in vivo two-photon calcium imaging of dendritic spines during visual stimulation and electron microscopy (EM) reconstructions of spine ultrastructure, investigating mitochondrial abundance near functionally- and structurally-characterized spines. Surprisingly, we found no correlation to structural measures of synaptic strength. Instead, we found that mitochondria are positioned near spines with orientation preferences that are dissimilar to the somatic preference. Additionally, we found that mitochondria are positioned near groups of spines with heterogeneous orientation preferences. For a subset of spines with mitochondrion in the head or neck, synapses were larger and exhibited greater selectivity to visual stimuli than those without a mitochondrion. Our data suggest mitochondria are not necessarily positioned to support the energy needs of strong spines, but rather support the structurally and functionally diverse inputs innervating the basal dendrites of cortical neurons.

neuroscience↗

A targeted, low-throughput compound screen in a Drosophila model of Neurofibromatosis Type 1 identifies simvastatin and BMS-204352 as potential therapies for autism spectrum disorder (ASD).

Autism spectrum disorder (ASD) is a common neurodevelopmental condition for which there are no pharmacological therapies that effectively target its core symptomatology. Animal models of syndromic forms of ASD, such as neurofibromatosis type 1, may be of use in screening for such treatments. Drosophila larvae lacking Nf1 expression exhibit tactile hypersensitivity following mechanical stimulation, proposed to mirror the sensory sensitivity issues comprising part of the ASD diagnostic criteria. Such behaviour is associated with synaptic dysfunction at the neuromuscular junction (NMJ). Both phenotypes may thus provide tractable outputs with which to screen for potential ASD therapies. In this study, we demonstrate that, while loss of Nf1 expression within the embryo is sufficient to impair NMJ synaptic transmission in the larva, constitutive Nf1 knockdown is required to induce tactile hypersensitivity, suggesting that a compound must be administered throughout development to rescue this behaviour. With such a feeding regime, we identify two compounds from a targeted, low-throughput screen that significantly and consistently reduce, but do not fully rescue, tactile hypersensitivity in Nf1P1 larvae. These are the HMG-CoA reductase inhibitor simvastatin, and the BKCa channel activator BMS-204352. At the NMJ, both compounds induce a significant reduction in the enhanced spontaneous transmission frequency of Nf1P1 larvae, though again not to the level of vehicle-treated controls. However, both compounds fully rescue the increased quantal size of Nf1P1 mutants, with simvastatin also fully rescuing their reduced quantal content. Thus, the further study of both compounds as potential ASD interventions is warranted. Significance StatementNo therapies currently exist that consistently and effectively target the core symptoms of autism spectrum disorder (ASD), which include altered responses to sensory stimuli. Previously it was shown that Drosophila larvae lacking expression of ASD-associated Nf1 display a heightened response to a mechanical stimulus and increased neuronal excitability, likely due to excessive Ras activity. Here, out of a screen for compounds targeting such mechanisms, we identified simvastatin and BMS-204352 to reduce the likelihood of a response in Nf1-/- larvae following mechanical stimulation. These compounds also improved synaptic transmission defects at the neuromuscular junction. Such findings support the further study of these drugs as potential ASD therapies in the clinic.

neuroscience↗

Cell-Type-Specific Epigenetic Priming of Gene Expression in Nucleus Accumbens by Cocaine

A hallmark of addiction is the ability of drugs of abuse to trigger relapse after periods of prolonged abstinence. Here, we describe a novel epigenetic mechanism whereby chronic cocaine exposure causes lasting chromatin and downstream transcriptional modifications in the nucleus accumbens (NAc), a critical brain region controlling motivation. We link prolonged withdrawal from cocaine to the depletion of the histone variant H2A.Z, coupled to increased genome accessibility and latent priming of gene transcription, in D1 dopamine receptor-expressing medium spiny neurons (D1 MSNs) that relates to aberrant gene expression upon drug relapse. The histone chaperone ANP32E removes H2A.Z from chromatin, and we demonstrate that D1 MSN-selective Anp32e knockdown prevents cocaine-induced H2A.Z depletion and blocks cocaines rewarding actions. By contrast, very different effects of cocaine exposure, withdrawal, and relapse were found for D2-MSNs. These findings establish histone variant exchange as an important mechanism and clinical target engaged by drugs of abuse to corrupt brain function and behavior.

molecular biology↗

Mitochondrial COA7 is a heme-binding protein involved in the early stages of complex IV assembly.

Cytochrome c oxidase assembly factor 7 (COA7) is a metazoan-specific assembly factor, critical for the biogenesis of mitochondrial complex IV (cytochrome c oxidase). Although mutations in COA7 have been linked in patients to complex IV assembly defects and neurological conditions such as peripheral neuropathy, ataxia and leukoencephalopathy, the precise role COA7 plays in the biogenesis of complex IV is not known. Here we show that the absence of COA7 leads to arrest of the complex IV assembly pathway at the initial step where the COX1 module is built, which requires incorporation of copper and heme cofactors. In solution, purified COA7 binds heme with micromolar affinity, through axial ligation to the central iron atom by histidine and methionine residues. Surprisingly, the crystal structure of COA7, determined to 2.4 [A] resolution, reveals a banana-shaped molecule composed of five helix-turn-helix (/) repeats, tethered by disulfide bonds, with a structure entirely distinct from proteins with characterized heme binding activities. We therefore propose a role for COA7 in heme binding/chaperoning in the mitochondrial intermembrane space, this activity being crucial for and providing a missing link in complex IV biogenesis. Significance StatementAssembly factors play key roles in the biogenesis of many mitochondrial protein complexes regulating their stability, activity and incorporation of essential cofactors. COA7 is a metazoan-specific assembly factor, the absence or mutation of which in humans accompanies complex IV assembly defects and neurological conditions. Here we report the crystal structure of COA7 to 2.4 [A] resolution, revealing a banana-shaped molecule composed of five helix-turn-helix (/) repeats, tethered by disulfide bonds. Characterization of pathogenic variants reveals significantly lower stabilities, correlating with the associated disease outcomes. Fascinatingly, COA7 binds heme with micromolar affinity, despite the fact that the protein structure does not resemble previously characterized heme-binding proteins. This provides a possible missing link for heme handling in the mitochondrial intermembrane space.

biochemistry↗

Mako: a graph-based pattern growth approach to detect complex structural variants

Complex structural variants (CSVs) are genomic alterations that have more than two breakpoints and are considered as simultaneous occurrence of simple structural variants. However, detecting the compounded mutational signals of CSVs is challenging through a commonly used model-match strategy. As a result, there has been limited progress for CSV discovery compared with simple structural variants. We systematically analyzed the multi-breakpoint connection feature of CSVs, and proposed Mako, utilizing a bottom-up guided model-free strategy, to detect CSVs from paired-end short-read sequencing. Specifically, we implemented a graph-based pattern growth approach, where the graph depicts potential breakpoint connections and pattern growth enables CSV detection without predefined models. Comprehensive evaluations on both simulated and real datasets revealed that Mako outperformed other algorithms. Notably, validation rates of CSV on real data based on experimental and computational validations as well as manual inspections are around 70%, where the medians of experimental and computational breakpoint shift are 13bp and 26bp, respectively. Moreover, Mako CSV subgraph effectively characterized the breakpoint connections of a CSV event and uncovered a total of 15 CSV types, including two novel types of adjacent segments swap and tandem dispersed duplication. Further analysis of these CSVs also revealed impact of sequence homology in the formation of CSVs. Mako is publicly available at https://github.com/jiadong324/Mako.

bioinformatics↗

Historical genomics reveals the evolutionary mechanisms behind multiple outbreaks of the coffee wilt pathogen Fusarium xylarioide

1Nearly 50% of crop yields are lost to pests and disease, with plants and pathogens locked in an amplified co-evolutionary process of disease outbreaks. Coffee wilt disease, caused by Fusarium xylarioides, decimated coffee production in west and central Africa following an initial 1920s outbreak. After successful management, it later re-emerged reaching two separate epidemics by the 2000s on arabica coffee in Ethiopia and robusta coffee in east and central Africa. Here, we use genome sequencing of six historical culture collection strains spanning 70 years to identify the evolutionary processes behind these repeated outbreaks. The robusta population arose from the initial outbreak, whilst the arabica population is divergent and emerged independently. The two populations evolved similar pathologies by separately acquiring different effector genes horizontally via transposable elements from other Fusarium taxa, including F. oxysporum. Thus, historical genomics can help reveal mechanisms that allow fungal pathogens to keep pace with humanitys efforts to resist them.

genomics↗

Cavin3 released from caveolae interacts with BRCA1to regulate the cellular stress response

Caveolae-associated protein 3 (cavin3), a putative tumor suppressor protein, is inactivated in most cancers. We characterized how cavin3 affects the cellular proteome using genome-edited cells together with label-free quantitative proteomics. These studies revealed a prominent role for cavin3 in DNA repair with BRCA1 and BRCA1 A-complex components being downregulated on cavin3 deletion. Cellular and cell-free expression assays, we show a direct interaction between BRCA1 and cavin3. Association of BRCA1 and cavin3 occurs when cavin3 is released from caveolae that are disassembled in response to UV and mechanical stress. Supporting a role in DNA repair, cavin3-deficient cells were sensitized to the effects of PARP inhibition, which compromises DNA repair, and showed reduced recruitment of the BRCA1 A-complex to UV DNA damage foci. Overexpression and RNAi-depletion revealed that cavin3 sensitized various cancer cells to UV-induced apoptosis. We conclude that cavin3 functions together with BRCA1 in multiple pathways that contribute to tumorigenesis.

cancer biology↗

Cortical neuron response selectivity derives from strength in numbers of synapses

Single neocortical neurons are driven by populations of excitatory inputs, forming the basis of neural selectivity to features of sensory input. Excitatory connections are thought to mature during development through activity-dependent Hebbian plasticity1, whereby similarity between presynaptic and postsynaptic activity selectively strengthens some synapses and weakens others2. Evidence in support of this process ranges from measurements of synaptic ultrastructure to slice and in vivo physiology and imaging studies3,4,5,6,7,8. These corroborating lines of evidence lead to the prediction that a small number of strong synaptic inputs drive neural selectivity, while weak synaptic inputs are less correlated with the functional properties of somatic output and act to modulate activity overall6,7. Supporting evidence from cortical circuits, however, has been limited to measurements of neighboring, connected cell pairs, raising the question of whether this prediction holds for the full profile of synapses converging onto cortical neurons. Here we measure the strengths of functionally characterized excitatory inputs contacting single pyramidal neurons in ferret primary visual cortex (V1) by combining in vivo two-photon synaptic imaging and post hoc electron microscopy (EM). Using EM reconstruction of individual synapses as a metric of strength, we find no evidence that strong synapses play a predominant role in the selectivity of cortical neuron responses to visual stimuli. Instead, selectivity appears to arise from the total number of synapses activated by different stimuli. Moreover, spatial clustering of co-active inputs, thought to amplify synaptic drive, appears reserved for weaker synapses, further enhancing the contribution of the large number of weak synapses to somatic response. Our results challenge the role of Hebbian mechanisms in shaping neuronal selectivity in cortical circuits, and suggest that selectivity reflects the co-activation of large populations of presynaptic neurons with similar properties and a mixture of strengths.

neuroscience↗

OpenCRAVAT, an open source collaborative platform for the annotation of human genetic variation

PURPOSEThe modern researcher is confronted with hundreds of published methods to interpret genetic variants. There are databases of genes and variants, phenotype-genotype relationships, algorithms that score and rank genes, and in silico variant effect prediction tools. Because variant prioritization is a multi-factorial problem, a welcome development in the field has been the emergence of decision support frameworks, which make it easier to integrate multiple resources in an interactive environment. Current decision support frameworks are typically limited by closed proprietary architectures, access to a restricted set of tools, lack of customizability, web dependencies that expose protected data, or limited scalability.\n\nMETHODSWe present OpenCRAVAT, a new open source, scalable decision support system for variant and gene prioritization. We have designed the resource catalog to be open and modular to maximize community and developer involvement, and as a result the catalog is being actively developed and growing every month. Resources made available via the store are well-suited for analysis of cancer, as well as Mendelian and complex diseases.\n\nRESULTSOpenCRAVAT offers both command line utility and dynamic GUI, allowing users to install with a single command, easily download tools from an extensive resource catalog, create customized pipelines, and explore results in a richly detailed viewing environment. We present several case studies to illustrate the design of custom workflows to prioritize genes and variants.\n\nCONCLUSIONOpenCRAVAT is distinguished from similar tools by its capabilities to access and integrate an unprecedented amount of diverse data resources and computational prediction methods, which span germline, somatic, common, rare, coding and non-coding variants. OpenCRAVAT is freely available at https://opencravat.org

bioinformatics↗

Plant respiration: controlled by photosynthesis or biomass?

Two simplifying hypotheses have been proposed for whole-plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first-principles carbon balance model with a prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carryover of fixed carbon between years, while the second implies far too great an increase in respiration during stand development - leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration is not linearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.

ecology↗