Forest Sciences Commons^™

Bacterium-Enabled Transient Gene Activation By Artificial Transcription Factor For Resolving Gene Regulation In Maize, Mingxia Zhao, Zhao Peng, Yang Qin, Ling Zhang, Bin Tian, Yueying Chen, Hairong Wei, Et. Al.

Michigan Tech Publications, Part 1

Cellular functions are diversified through intricate transcription regulations, and an understanding gene regulation networks is essential to elucidating many developmental processes and environmental responses. Here, we employed the Transcriptional-Activator Like effectors (TALes), which represent a family of transcription factors that are synthesized by members of the γ-proteobacterium genus Xanthomonas and secreted to host cells for activation of targeted host genes. Through delivery by the maize pathogen, Xanthomonas vasicola pv. vasculorum, designer TALes (dTALes), which are synthetic TALes, were used to induce the expression of the maize gene glossy3 (gl3), a MYB transcription factor gene involved in the …

Microrna319-Mediated Gene Regulatory Network Impacts Leaf Development And Morphogenesis In Poplar, Yanxia Cheng, Lihu Wang, Manzar Abbas, Xiong Huang, Qiao Wang, Aimin Wu, Hairong Wei, Et. Al.

Michigan Tech Publications, Part 1

MicroRNA319 (miR319) has been implicated in leaf development in a number of plant species. Here we study the roles of miR319a and its regulated network in leaf development in poplars. Over-expression of miR319a in Populus alba × Populus glandulosa caused dwarf statures, narrow leaf blades and serrated leaf margins. The vascular bundles and bundle sheaths in transgenic leaves had more layers of cells than those in the leaves of control plants, indicating enhanced lignification in these cells. Among the 93 putative targets of miR319a predicted with the psRNATarget tool, only three genes, TCP (TEOSINTE BRANCHED1, CYCLOIDEA, and …

Consequences Of Increased Variation In Peatland Hydrology For Carbon Storage: Legacy Effects Of Drought And Flood In A Boreal Fen Ecosystem, Evan Kane, Catherine M. Dieleman, Danielle L. Rupp, Kevin H. Wyatt

Michigan Tech Publications, Part 1

Globally important carbon (C) stores in boreal peatlands are vulnerable to altered hydrology through changes in precipitation and runoff patterns, groundwater inputs, and a changing cryosphere. These changes can affect the extent of boreal wetlands and their ability to sequester and transform C and other nutrients. Variation in precipitation patterns has also been increasing, with greater occurrences of both flooding and drought periods. Recent work has pointed to the increasing role of algal production in regulating C cycling during flooded periods in fen peatlands, but exactly how this affects the C sink-strength of these ecosystems is poorly understood. We evaluated …

Inaugural Editorial, Hairong Wei

Michigan Tech Publications, Part 1

No abstract provided.

Growth-Regulating Factor 5 (Grf5)-Mediated Gene Regulatory Network Promotes Leaf Growth And Expansion In Poplar, Wenqi Wu, Jiang Li, Qiao Wang, Kaiwen Lv, Kang Du, Hairong Wei

Michigan Tech Publications, Part 1

Although polyploid plants have larger leaves than their diploid counterparts, the molecular mechanisms underlying this difference (or trait) remain elusive. Differentially expressed genes (DEGs) between triploid and full-sib diploid poplar trees were identified from two transcriptomic data sets followed by a gene association study among DEGs to identify key leaf growth regulators. Yeast one-hybrid system, electrophoretic mobility shift assay, and dual-luciferase assay were employed to substantiate that PpnGRF5-1 directly regulated PpnCKX1. The interactions between PpnGRF5-1 and growth-regulating factor (GRF)-interacting factors (GIFs) were experimentally validated and a multilayered hierarchical regulatory network (ML-hGRN)-mediated by PpnGRF5-1 was constructed with top-down graphic Gaussian model …

Fine-Scale Movements And Behaviors Of Coyotes (Canis Latrans) During Their Reproductive Period, Michael J. Chamberlain, Bradley S. Cohen, Patrick H. Wightman, Emily Rushton, Joseph W. Hinton

Michigan Tech Publications, Part 1

In canids, resident breeders hold territories but require different resources than transient individuals (i.e., dispersers), which may result in differential use of space, land cover, and food by residents and transients. In the southeastern United States, coyote (Canis latrans) reproduction occurs during spring and is energetically demanding for residents, but transients do not reproduce and therefore can exhibit feeding behaviors with lower energetic rewards. Hence, how coyotes behave in their environment likely differs between resident and transient coyotes. We captured and monitored 36 coyotes in Georgia during 2018–2019 and used data from 11 resident breeders, 12 predispersing residents (i.e., offspring …

Root Biomass And Production By Two Cushion Plant Species Of Tropical High-Elevation Peatlands In The Andean Páramo, Esteban Suárez, Segundo Chimbolema, Rodney Chimner, Erik A. Lilleskov

Michigan Tech Publications, Part 1

High-elevation peatlands in the Andes are receiving increasing attention for their biodiversity and their high rates of carbon accumulation. However, the ecology of these peatlands and the environmental factors that control their carbon dynamics remain under-studied. Here we report on the patterns of root biomass productivity and turnover rates for two cushion plant species (Distichia muscoides, Plantago rigida) that commonly dominate high-elevation peatlands (> 4200 m a.s.l.) in the Andean páramo landscape of Northern Ecuador. Root biomass for P. rigida ranged from 680 to 864 g m-2 and was approximately 40 % higher than for D. muscoides (507–620 g m-2). …

High-Quality De Novo Assembly Of The Eucommia Ulmoides Haploid Genome Provides New Insights Into Evolution And Rubber Biosynthesis, Yun Li, Hairong Wei, Jun Yang, Kang Du, Jiang Li, Ying Zhang, Tong Qiu, Zhao Liu, Yongyu Ren, Lianjun Song, Xiangyang Kang

Michigan Tech Publications, Part 1

We report the acquisition of a high-quality haploid chromosome-scale genome assembly for the first time in a tree species, Eucommia ulmoides, which is known for its rubber biosynthesis and medicinal applications. The assembly was obtained by applying PacBio and Hi–C technologies to a haploid that we specifically generated. Compared to the initial genome release, this one has significantly improved assembly quality. The scaffold N50 (53.15 MB) increased 28-fold, and the repetitive sequence content (520 Mb) increased by 158.24 Mb, whereas the number of gaps decreased from 104,772 to 128. A total of 92.87% of the 26,001 predicted protein-coding genes identified …

High-Purity Recycling Of Hematite And Zn/Cu Mixture From Waste Smelting Slag, Yang Huo, Xiang Song, Suiyi Zhu, Yu Chen, Xue Lin, Yaqiong Wu, Zhan Qu, Ting Su, Xinfeng Xie

Michigan Tech Publications, Part 1

In this study, Zn/Cu-bearing smelting slag was recycled via an integrated acid dissolution and hematite precipitation method. The slag was dissolved in nitric acid to generate an acid solution containing 23.5 g/L Fe, 4.45 g/L Zn and 2.81 g/L Cu, which was subjected to hydrothermal treatment with the addition of levulinic acid (LA). More than 99.95% of the initial Fe content was removed as hematite particles with diameters of approximately 200 nm, and the residual Fe concentration in the acid was 0.43 mg/L. The generated hematite contained 97.3% Fe2O3, 0.64% ZnO and 0.58% CuO. Greater than 99% of the initial …

Comparative Proteomic Analysis Of Leaves At Different Ages In Allotriploid Populus, Jiang Li, Yi Wang, Hairong Wei, Xiangyang Kang

Michigan Tech Publications, Part 1

Triploid poplar trees have been shown to have a number of growth advantages, especially much bigger leaves that contribute greatly to the increased biomass. In this study, we focused on the relationships between leaf age and leaf metabolism in triploids. We performed comparative proteomic analysis of the 5th (FDR5), 10th (FDR10), and 25th (FDR25) leaves from the apical meristems in allotriploids originated from first-division restitution (FDR). A total of 1970, 1916, and 1850 proteins were identified in the FDR5, FDR10, and FDR25, respectively. Principle component analysis (PCA) and differentially accumulated protein (DAP) analysis showed that FDR10 and FDR25 displayed higher …

Climate, Snowmelt Dynamics And Atmospheric Deposition Interact To Control Dissolved Organic Carbon Export From A Northern Forest Stream Over 26 Years, Karl M. Meingast, Evan Kane, Ashley A. Coble, Amy Marcarelli, David Toczydlowski

Michigan Tech Publications, Part 1

Increasing concentrations of dissolved organic carbon (DOC) have been identified in many freshwater systems over the last three decades. Studies have generally nominated atmospheric deposition as the key driver of this trend, with changes in climatic factors also contributing. However, there is still much uncertainty concerning net effects of these drivers on DOC concentrations and export dynamics. Changes in climate and climate mediated snowfall dynamics in northern latitudes have not been widely considered as causal factors of changes in long-term DOC trends, despite their disproportionate role in annual DOC export. We leveraged long-term datasets (1988–2013) from a first-order forested tributary …

Enrollment Decision-Making By Students In Forestry And Related Natural Resource Degree Programmes Globally, T. L. Bal, M. D. Rouleau, T. L. Sharik, A. M. Wellstead

Michigan Tech Publications, Part 1

A survey of 396 undergraduate and graduate students from 51 countries on 5 continents currently enrolled in Forestry or Related Natural Resource (FRNR) degree programmes was conducted of attendees to the International Union of Forest Research Organizations' (IUFRO) conference in Salt Lake City, Utah, 2014. These perspectives come from some of the most active students in their respective fields. We explored the motivating reasons for enrolling in their current FRNR programme, and conversely why they may have been hesitant to do so. Results indicate that enjoyment of nature was the most important factor on average driving the decision to enroll, …

Photosynthetic And Respiratory Acclimation Of Understory Shrubs In Response To In Situ Experimental Warming Of A Wet Tropical Forest, Kelsey R. Carter, Tana E. Wood, Sasha C. Reed, Elsa Schwartz, Madeline B. Reinsel, Xi Yang, Molly A. Cavaleri

Michigan Tech Publications, Part 1

Despite the importance of tropical forests to global carbon balance, our understanding of how tropical plant physiology will respond to climate warming is limited. In addition, the contribution of tropical forest understories to global carbon cycling is predicted to increase with rising temperatures, however, in situ warming studies of tropical forest plants to date focus only on upper canopies. We present results of an in situ field-scale +4°C understory infrared warming experiment in Puerto Rico (Tropical Responses to Altered Climate Experiment; TRACE). We investigated gas exchange responses of two common understory shrubs, Psychotria brachiata and Piper glabrescens, after exposure …

Vegetative And Edaphic Responses In A Northern Mixed Conifer Forest Three Decades After Harvest And Fire: Implications For Managing Regeneration And Carbon And Nitrogen Pools, R. Kasten Dumroese, Martin Jurgensen, Deborah S. Page-Dumroese

Michigan Tech Publications, Part 1

Research Highlights: This experiment compares a range of combinations of harvest, prescribed fire, and wildfire. Leveraging a 30-year-old forest management-driven experiment, we explored the recovery of woody species composition, regeneration of the charismatic forest tree species Larix occidentalis Nutt., and vegetation and soil carbon (C) and nitrogen (N) pools. Background and Objectives: Initiated in 1967, this experiment intended to explore combinations of habitat type phases and prescribed fire severity toward supporting regeneration of L. occidentalis. At onset of the experiment, a wildfire affected a portion of the 60 research plots, allowing for additional study. Our objective was to better understand …

Overexpression Of Bplerd15 Enhances Drought Tolerance In Betula Platyphylla Suk, Kaiwen Lv, Hairong Wei, Jing Jiang

Michigan Tech Publications, Part 1

In this study, we report the cloning and functional characterization of an early responsive gene, BplERD15, from Betula platyphylla Suk to dehydration. BplERD15 is located in the same branch as Morus indica Linnaeus ERD15 and Arabidopsis Heynh ERD15 in the phylogenetic tree built with ERD family protein sequences. The tissue-specific expression patterns of BplERD15 were characterized using qRT-PCR and the results showed that the transcript levels of BplERD15 in six tissues were ranked from the highest to the lowest levels as the following: mature leaves (ML) > young leaves (YL) > roots (R) >buds (B) >young stems (YS) >mature stems (MS). Multiple …

Wetland And Hydric Soils, Carl C. Trettin, Randall K. Kolka, Anne S. Marsh, Sheel Bansal, Erik A. Lilleskov, Rodney Chimner, Et Al.

Michigan Tech Publications, Part 1

Soil and the inherent biogeochemical processes in wetlands contrast starkly with those in upland forests and rangelands. The differences stem from extended periods of anoxia, or the lack of oxygen in the soil, that characterize wetland soils; in contrast, upland soils are nearly always oxic. As a result, wetland soil biogeochemistry is characterized by anaerobic processes, and wetland vegetation exhibits specific adaptations to grow under these conditions. However, many wetlands may also have periods during the year where the soils are unsaturated and aerated. This fluctuation between aerated and nonaerated soil conditions, along with the specialized vegetation, gives rise to …

Comparing Human Activity Density And Green Space Supply Using The Baidu Heat Map In Zhengzhou, China, Shumei Zhang, Wenshi Zhang, Ying Wang, Xiaoyu Zhao, Peihao Song, Guohang Tian, Audrey L. Mayer

Michigan Tech Publications, Part 1

Rapidly growing cities often struggle with insufficient green space, although information on when and where more green space is needed can be difficult to collect. Big data on the density of individuals in cities collected from mobile phones can estimate the usage intensity of urban green space. Taking Zhengzhou's central city as an example, we combine the real-time human movement data provided by the Baidu Heat Map, which indicates the density of mobile phones, with vector overlays of different kinds of green space. We used the geographically weighted regression (GWR) method to estimate differentials in green space usage between weekdays …

Insights On Dissolved Organic Matter Production Revealed By Removal Of Charge-Transfer Interactions In Senescent Leaf Leachates, Karl M. Meingast, Brice K. Grunert, Sarah A. Green, Evan S. Kane, Nastaran Khademimoshgenani

Michigan Tech Publications, Part 1

Dissolved organic matter (DOM) is a critical part of the global carbon cycle. Currently, it is understood that at least a portion of the chromophoric DOM (CDOM) character can be described through an electronic interaction of charge transfer (CT) complexes. While much work has been done to understand the influence of CT on soil and aquatic reference standard DOM, little is known about the influence of CT in fresh terrestrially derived DOM. In this study, leaf litter leachates from three tree species were treated (reduced) with sodium borohydride to determine the contribution of CT on a source of fresh terrestrial …

Tropical Understory Herbaceous Community Responds More Strongly To Hurricane Disturbance Than To Experimental Warming, Deborah K. Kennard, David Matlaga, Joanne Sharpe, Clay King, Aura M. Alonso-Rodríguez, Sasha C. Reed, Molly A. Cavaleri, Tana E. Wood

Michigan Tech Publications, Part 1

Ecology and Evolution published by John Wiley & Sons Ltd. The effects of climate change on tropical forests may have global consequences due to the forests’ high biodiversity and major role in the global carbon cycle. In this study, we document the effects of experimental warming on the abundance and composition of a tropical forest floor herbaceous plant community in the Luquillo Experimental Forest, Puerto Rico. This study was conducted within Tropical Responses to Altered Climate Experiment (TRACE) plots, which use infrared heaters under free-air, open-field conditions, to warm understory vegetation and soils + 4°C above nearby control plots. Hurricanes …

Multiple Fgf4 Retrocopies Recently Derived Within Canids, Kevin Batcher, Peter Dickinson, Kimberly Maciejczyk, Kristin Brzeski, Sheida Hadji Rasouliha, Anna Letko, Cord Drögemüller, Tosso Leeb, Danika Bannasch

Michigan Tech Publications, Part 1

Two transcribed retrocopies of the fibroblast growth factor 4 (FGF4) gene have previously been described in the domestic dog. An FGF4 retrocopy on chr18 is associated with disproportionate dwarfism, while an FGF4 retrocopy on chr12 is associated with both disproportionate dwarfism and intervertebral disc disease (IVDD). In this study, whole-genome sequencing data were queried to identify other FGF4 retrocopies that could be contributing to phenotypic diversity in canids. Additionally, dogs with surgically confirmed IVDD were assayed for novel FGF4 retrocopies. Five additional and distinct FGF4 retrocopies were identified in canids including a copy unique to red wolves (Canis rufus). The …

Hydrological Effects Of Urban Green Space On Stormwater Runoff Reduction In Luohe, China, Peihao Song, Jianhui Guo, Enkai Xu, Audrey L. Mayer, Chang Liu, Jing Huang, Guohang Tian, Gunwoo Kim

Michigan Tech Publications, Part 1

This paper reveals the role of urban green space (UGS) in regulating runoff and hence on urban hydrological balance. The modeling software i-Tree Hydro was used to quantify the effects of UGS on surface runoff regulation and canopy interception capacity in four simulated land-cover scenarios. The results showed that the existing UGS could mitigate 15,871,900 m3 volume of runoff (accounting for 9.85% of total runoff) and intercept approximately 9.69% of total rainfall by the vegetation canopy. UGS in midterm goal and final goal scenarios could retain about 10.74% and 10.89% of total rainfall that falls onto the canopy layer, respectively. …

Wood Decomposition After An Aerial Application Of Hydromulch Following Wildfire In A Southern California Chaparral Shrubland, Martin Jurgensen, Chris Miller, Deborah S. Page-Dumroese

Michigan Tech Publications, Part 1

Severe wildfire can affect many soil processes, especially organic matter (OM) decomposition. Organic mulches are often applied on steep slopes to mitigate soil erosion, but little is known about how these surface organic additions affect subsequent soil OM decomposition. In 2003 the Cedar Fire burned 110,000 ha in southern California chaparral shrubland, after which hydromulch was aerially applied to reduce soil erosion. Subsequently, we established a 5-year study to assess the effect of hydromulch on OM decomposition at the burned soil surface and in the mineral soil using aspen (Populus tremuloides Michx.) and pine (Pinus taeda L.) wood stakes as …

Patterns Of Ecosystem Structure And Wildfire Carbon Combustion Across Six Ecoregions Of The North American Boreal Forest, Xanthe J. Walker, Jennifer L. Baltzer, Laura Bourgeau-Chavez, Nicola J. Day, Catherine M. Dieleman, Jill F. Johnstone, Evan Kane, Brendan M. Rogers, Merritt R. Turetsky, Sander Veraverbeke, Michelle C. Mack

Michigan Tech Publications, Part 1

Increases in fire frequency, extent, and severity are expected to strongly impact the structure and function of boreal forest ecosystems. An important function of the boreal forest is its ability to sequester and store carbon (C). Increasing disturbance from wildfires, emitting large amounts of C to the atmosphere, may create a positive feedback to climate warming. Variation in ecosystem structure and function throughout the boreal forest is important for predicting the effects of climate warming and changing fire regimes on C dynamics. In this study, we compiled data on soil characteristics, stand structure, pre-fire C pools, C loss from fire, …

Spatio-Temporal Dynamics Of Landscape Use By The Bumblebee Bombus Pauloensis (Hymenoptera: Apidae) And Its Relationship With Pollen Provisioning, Pablo Cavigliasso, Colin Phifer, Erika M. Adams, David J. Flaspohler, Gerardo P. Gennari, Julian A. Licata, Natacha P. Chacoff

Michigan Tech Publications, Part 1

Understanding how bees use resources at a landscape scale is essential for developing meaningful management plans that sustain populations and the pollination services they provide. Bumblebees are important pollinators for many wild and cultivated plants, and have experienced steep population declines worldwide. Bee foraging behavior can be influenced by resource availability and bees' lifecycle stage. To better understand these relationships, we studied the habitat selection of Bombus pauloensis by tracking 17 queen bumblebees with radio telemetry in blueberry fields in Entre Ríos province, Argentina. To evaluate land use and floral resources used by bumblebees, we tracked bees before and after …

Predicting Effects Of Climate Change On Productivity And Persistence Of Forest Trees, Russell D. Kramer, H. Roaki Ishii, Kelsey R. Carter, Yuko Miyazaki, Molly A. Cavaleri, Masatake G. Araki, Wakana A. Azuma, Yuta Inoue, Chinatsu Hara

Michigan Tech Publications, Part 1

Global climate change increases uncertainty in sustained functioning of forest ecosystems. Forest canopies are a key link between terrestrial ecosystems, the atmosphere, and climate. Here, we introduce research presented at the 66th meeting of the Ecological Society of Japan in the symposium “Structure and function of forest canopies under climate change.” Old-growth forest carbon stores are the largest and may be the most vulnerable to climate change as the balance between sequestration and emission could easily be tipped. Detailed structural analysis of individual large, old trees shows they are allocating wood to the trunk and crown in patterns that cannot …

Taxonomic Similarity Does Not Predict Necessary Sample Size For Ex Situ Conservation: A Comparison Among Five Genera, Sean Hoban, Taylor Callicrate, John Clark, Susan Deans, Michael Dosmann, Priyanka Dipak Kadav, Et Al.

Michigan Tech Publications, Part 1

Effectively conserving biodiversity with limited resources requires scientifically informed and efficient strategies. Guidance is particularly needed on how many living plants are necessary to conserve a threshold level of genetic diversity in ex situ collections. We investigated this question for 11 taxa across five genera. In this first study analysing and optimizing ex situ genetic diversity across multiple genera, we found that the percentage of extant genetic diversity currently conserved varies among taxa from 40% to 95%. Most taxa are well below genetic conservation targets. Resampling datasets showed that ideal collection sizes vary widely even within a genus: one taxon …

Conceptual Planning Of Urban–Rural Green Space From A Multidimensional Perspective: A Case Study Of Zhengzhou, China, Bo Mu, Chang Liu, Guohang Tian, Yaqiong Xu, Yali Zhang, Audrey L. Mayer, Et Al.

Michigan Tech Publications, Part 1

The structure and function of green-space system is an eternal subject of landscape architecture, especially due to limited land and a need for the coordinated development of PLEs (production, living, and ecological spaces). To make planning more scientific, this paper explored green-space structure planning via multidimensional perspectives and methods using a case study of Zhengzhou. The paper applies theories (from landscape architecture and landscape ecology) and technologies (like remote sensing, GIS—geographic information system, graph theory, and aerography) from different disciplines to analyze current green-space structure and relevant physical factors to identify and exemplify different green-space planning strategies. Overall, our analysis …

Trends In Protected Area Representation Of Biodiversity And Ecosystem Services In Five Tropical Countries, Rachel A. Neugarten, Kevin Moull, Natalia Acero Martinez, Luciano Andriamaro, Curtis Bernard, Leonardo Saenz, Et Al.

Michigan Tech Publications, Part 1

In late 2020, governments will set the next decade of conservation targets under the UN Convention on Biological Diversity. Setting new targets requires understanding how well national protected area (PA) networks are spatially representing important areas for biodiversity and ecosystem services. We analyzed the representation of biodiversity priority areas (BPAs), forests, forest carbon stocks, non-timber forest products (NTFPs), and freshwater ecosystem services (FES) within terrestrial PA systems in Cambodia, Guyana, Liberia, Madagascar, and Suriname in 2003 and 2017. Four of the countries (all except Suriname) expanded their terrestrial PA networks during the study period. In all five countries, we found …

Effects Of A Regenerating Matrix On The Survival Of Birds In Tropical Forest Fragments, Jared D. Wolfe, Philip C. Stouffer, Richard O. Bierregaard Jr., David A, Luther, Thomas E. Lovejoy

Michigan Tech Publications, Part 1

Background

Vast areas of lowland neotropical forest have regenerated after initially being cleared for agricultural purposes. The ecological value of regenerating second growth to forest-dwelling birds may largely depend on the age of the forest, associated vegetative structure, and when it is capable of sustaining avian demographics similar to those found in pristine forest.

Methods

To determine the influence of second growth age on bird demography, we estimated the annual survival of six central Amazonian bird species residing in pristine forest, a single 100 and a single 10 ha forest fragment, taking into consideration age of the surrounding matrix (i.e. …

A Phylogenomic Approach Reveals A Low Somatic Mutation Rate In A Long-Lived Plant., Adam J Orr, Amanda Padovan, David Kainer, Carsten Kulheim, Lindell Bromham, Carlos Bustos-Segura, William Foley, Tonya Haff, Ji-Fan Hsieh, Alejandro Morales-Suarez, Reed A Cartwright, Robert Lanfear

Michigan Tech Publications, Part 1

Somatic mutations can have important effects on the life history, ecology, and evolution of plants, but the rate at which they accumulate is poorly understood and difficult to measure directly. Here, we develop a method to measure somatic mutations in individual plants and use it to estimate the somatic mutation rate in a large, long-lived, phenotypically mosaic Eucalyptus melliodora tree. Despite being 100 times larger than Arabidopsis, this tree has a per-generation mutation rate only ten times greater, which suggests that this species may have evolved mechanisms to reduce the mutation rate per unit of growth. This adds to a …