The bright side of parasitic plants: what are they good for?

Parasitic plants are mostly viewed as pests. This is caused by several species causing serious damage to agriculture and forestry. There is however much more to parasitic plants than presumed weeds. Many parasitic plans exert even positive effects on natural ecosystems and human society, which we review in this paper. Plant parasitism generally reduces the growth and fitness of the hosts. The network created by a parasitic plant attached to multiple host plant individuals may however trigger transferring systemic signals among these. Parasitic plants have repeatedly been documented to play the role of keystone species in the ecosystems. Harmful effects on community dominants, including invasive species, may facilitate species coexistence and thus increase biodiversity. Many parasitic plants enhance nutrient cycling and provide resources to other organisms like herbivores or pollinators, which contributes to facilitation cascades in the ecosystems. There is also a long tradition of human use of parasitic plants for medicinal and cultural purposes worldwide. Few species provide edible fruits. Several parasitic plants are even cultivated by agriculture/forestry for efficient harvesting of their products. Horticultural use of some parasitic plant species has also been considered. While providing multiple benefits, parasitic plants should always be used with care. In particular, parasitic plant species should not be cultivated outside their native geographical range to avoid the risk of their uncontrolled spread and the resulting damage to ecosystems.

Advances

• Parasitic plants may act as highways for transferring systemic signals among host plants.
• Harmful effects of parasitic plants on individual hosts suppress community dominants including invasive species, reduce competitive pressure, and may increase biodiversity.
• Parasitic plants enhance nutrient cycling and provide resources for other organisms thus contributing to facilitation cascades in ecosystems.
• Many parasitic plants are recorded to have medicinal values against a broad range of diseases.
• There is a long tradition of worldwide human use of parasitic plants, which have been cultivated for their products and aesthetic values.

     

Adaptation of the parasitic plant lifecycle: germination is controlled by essential host signaling molecules

Parasitic plants are plants that connect with a haustorium to the vasculature of another, host, plant from which they absorb water, assimilates, and nutrients. Because of this parasitic lifestyle, parasitic plants need to coordinate their lifecycle with that of their host. Parasitic plants have evolved a number of host detection/host response mechanisms of which the germination in response to chemical host signals in one of the major families of parasitic plants, the Orobanchaceae, is a striking example. In this update review, we discuss these germination stimulants. We review the different compound classes that function as germination stimulants, how they are produced, and in which host plants. We discuss why they are reliable signals, how parasitic plants have evolved mechanisms that detect and respond to them, and whether they play a role in host specificity. The advances in the knowledge underlying this signaling relationship between host and parasitic plant have greatly improved our understanding of the evolution of plant parasitism and are facilitating the development of more effective control measures in cases where these parasitic plants have developed into weeds.

Root parasitic plants grow on the roots of other plants and germinate only in the presence of that host, on which they completely depend, through the perception of host presence signaling molecules called germination stimulants.

Outstanding questions

• Have we overlooked the role of germination stimulants in facultative parasites?
• What is the biological relevance of the observation that many plant species produce and secrete a range of different strigolactones?
• Have parasitic plants evolved mechanisms to compensate for low phosphorus availability, a condition that stimulates their germination?
• What is the contribution of the HTL strigolactone receptors to host specificity in parasitic plants or does downstream signaling play a role?
• What other, nonstrigolactone, germination stimulants can parasitic plants respond to and does this require adaptation in the HTL receptors?
• What is the role of germination and underlying mechanism in the rapid adaptation of (orobanchaceous) parasitic plants to a new host?

     

The mechanism of host-induced germination in root parasitic plants

Chemical signals known as strigolactones (SLs) were discovered more than 50 years ago as host-derived germination stimulants of parasitic plants in the Orobanchaceae. Strigolactone-responsive germination is an essential adaptation of obligate parasites in this family, which depend upon a host for survival. Several species of obligate parasites, including witchweeds (Striga, Alectra spp.) and broomrapes (Orobanche, Phelipanche spp.), are highly destructive agricultural weeds that pose a significant threat to global food security. Understanding how parasites sense SLs and other host-derived stimulants will catalyze the development of innovative chemical and biological control methods. This review synthesizes the recent discoveries of strigolactone receptors in parasitic Orobanchaceae, their signaling mechanism, and key steps in their evolution.

A family of receptors that evolved in the Orobanchaceae family enable seeds of parasitic plants to sense strigolactones from a nearby host root and germinate.

Advances

• Strigolactone perception by parasite seed is mediated by a clade of neofunctionalized KAI2d proteins that evolved from a receptor that mediates karrikin responses in other plants.
• KAI2d proteins use a similar mechanism to perceive SLs as D14, which mediates growth responses to SLs in nonparasites, but activate different signaling pathways.
• Crystal structure analyses and chemical probes reveal features of KAI2d ligand-binding pockets that contribute to their specificity.

     

Holoparasitic plant–host interactions and their impact on Mediterranean ecosystems

Although photosynthesis is essential to sustain life on Earth, not all plants use sunlight to synthesize nutrients from carbon dioxide and water. Holoparasitic plants, which are important in agricultural and natural ecosystems, are dependent on other plants for nutrients. Phytohormones are crucial in holoparasitic plant–host interactions, from seed germination to senescence, not only because they act as growth and developmental regulators, but also because of their central role in the regulation of host photosynthesis and source–sink relations between the host and the holoparasitic plant. Here, we compile and discuss current knowledge on the impact and ecophysiology of holoparasitic plants (such as the broomrapes Orobanche sp. and Phelipanche sp.) that infest economically important dicotyledonous crops in Mediterranean agroecosystems (legumes [Fabaceae], sunflowers [Helianthus sp.], or tomato [Solanum lycopersicum] plants). We also highlight the role of holoparasitic plant–host interactions (such as those between Cytinus hypocistis and various shrubs of the genus Cistus) in shaping natural Mediterranean ecosystems. The roles of phytohormones in controlling plant–host interactions, abiotic factors in parasitism, and the biological significance of natural seed banks and how dormancy and germination are regulated, will all be discussed. Holoparasitic plants are unique organisms; improving our understanding of their interaction with hosts as study models will help us to better manage parasitic plants, both in agricultural and natural ecosystems.

Advances

• Mediterranean ecosystems represent unique environments to study holoparasitic plant-host interactions
• Holoparasitic plants cause severe reductions in productivity, but can also exert positive effects on diversity in natural ecosystems
• A bidirectional flux of phytohormones occurs in holoparasitic plant-host interactions
• The establishment of seed banks is essential for the success of both Orobanche and Cytinus infection in Mediterranean ecosystems

     

Live monitoring of plant redox and energy physiology with genetically encoded biosensors

Genetically encoded biosensors pave the way for understanding plant redox dynamics and energy metabolism on cellular and subcellular levels.

ADVANCES

• Methodological advances in fluorescent protein-based in vivo biosensing have been instrumental for several paradigm shifts in our understanding of cell physiology, metabolism and signaling.
• An increasing number of genetically encoded biosensors has been used to dissect the dynamics of several distinct redox couples and energy physiology in plants.
• In vivo monitoring using biosensors has pioneered the simultaneous read-out of different physiological parameters in different subcellular locations by parallelized plate reader-based, multiwell fluorimetry, or expression strategies for multiple sensors in parallel.
• Sensing dynamic changes in hydrogen peroxide levels is possible with sensors of the HyPer family, or roGFP fusion variants with a thiol peroxidase.
• Peredox and SoNar family sensors enable direct visualization of NADH/NAD⁺, while iNAP family sensors respond to NADPH concentration in plants.
• Sensor variants with different sensitivity ranges enable use of the most appropriate variant for the specific in vivo environment or experimental scope.

     

Contemporary proteomic strategies for cysteine redoxome profiling

Protein cysteine residues are susceptible to oxidative modifications that can affect protein functions. Proteomic techniques that comprehensively profile the cysteine redoxome, the repertoire of oxidized cysteine residues, are pivotal towards a better understanding of the protein redox signaling. Recent technical advances in chemical tools and redox proteomic strategies have greatly improved selectivity, in vivo applicability, and quantification of the cysteine redoxome. Despite this substantial progress, still many challenges remain. Here, we provide an update on the recent advances in proteomic strategies for cysteine redoxome profiling, compare the advantages and disadvantages of current methods and discuss the outstanding challenges and future perspectives for plant redoxome research.

Current cysteine redoxome profiling can characterize systematically diverse oxidative posttranslational modifications

Advances

• The chemical toolbox for Cys redoxome profiling has extensively expanded.
• Advanced chemoproteomic platforms have been applied to target specific Cys oxidative posttranslational modifications (OxiPTMs).
• Various reductomic workflows have been widely implemented for reversible Cys OxiPTMs quantification.
• Workflows have been integrated to measure the occupancy of multiple OxiPTMs simultaneously.
• Disulfide-based traps enable the in situ profiling for –SOH sites.

     

植物寄生对生态系统结构和功能的影响

寄生植物是生态系统中的特殊类群之一。植物寄生可以驱动生态系统中生物与非生物因子的变化,在生态系统结构与功能中起关键作用。寄生植物可以通过对寄主营养的集聚、改变凋落物的质量与数量、改变根的周转与分泌物格局、改变土壤水势,从而影响土壤理化特性。寄生植物会改变寄主的行为,改变寄主与非寄主植物之间的相互作用,从而影响植物群落的结构、多样性和动态,进而影响植被演替和植被生产力等。寄生植物与寄主均可被消费者取食,可直接或间接地影响生态系统的食草动物,包括草食昆虫等。寄生植物与寄主的其它寄生物存在竞争关系,可以直接或间接地影响寄主的其它寄生植物或病原真菌。寄生植物可以明显地改变土壤地球化学循环,将固有的不可动的成分转变为可利用的营养成分,改变土壤生物群落的结构与功能,从而显著影响地下生物群落。这些表明,植物寄生对生态系统的结构和功能有重要影响。针对特殊的被入侵的植物群落,该地寄生植物可以通过影响入侵植物寄主的生长、繁殖、生物量分配格局,改变土壤的理化特性,促进非寄主的非优势本地植物的生长,从而改变被入侵植物群落结构与多样性,达到生物防治及生态恢复的目的。     

Subtilase activity in intrusive cells mediates haustorium maturation in parasitic plants

Parasitic plants that infect crops are devastating to agriculture throughout the world. These parasites develop a unique inducible organ called the haustorium that connects the vascular systems of the parasite and host to establish a flow of water and nutrients. Upon contact with the host, the haustorial epidermal cells at the interface with the host differentiate into specific cells called intrusive cells that grow endophytically toward the host vasculature. Following this, some of the intrusive cells re-differentiate to form a xylem bridge (XB) that connects the vasculatures of the parasite and host. Despite the prominent role of intrusive cells in host infection, the molecular mechanisms mediating parasitism in the intrusive cells remain poorly understood. In this study, we investigated differential gene expression in the intrusive cells of the facultative parasite Phtheirospermum japonicum in the family Orobanchaceae by RNA-sequencing of laser-microdissected haustoria. We then used promoter analyses to identify genes that are specifically induced in intrusive cells, and promoter fusions with genes encoding fluorescent proteins to develop intrusive cell-specific markers. Four of the identified intrusive cell-specific genes encode subtilisin-like serine proteases (SBTs), whose biological functions in parasitic plants are unknown. Expression of SBT inhibitors in intrusive cells inhibited both intrusive cell and XB development and reduced auxin response levels adjacent to the area of XB development. Therefore, we propose that subtilase activity plays an important role in haustorium development in P. japonicum.

Subtilases specifically expressed in intrusive cells regulate auxin-mediated host–parasite connections in the parasitic plant Phtheirospermum japonicum.     

Specific developmental pathways underlie host specificity in the parasitic plant Orobanche

Parasitic angiosperms are an ecologically and economically important group of plants. However our understanding of the basis for host specificity in these plants is embryonic. Recently we investigated host specificity in the parasitic angiosperm Orobanche minor, and demonstrated that this host generalist parasite comprises genetically defined races that are physiologically adapted to specific hosts. Populations occurring naturally on red clover (Trifolium pratense) and sea carrot (Daucus carota subsp. gummifer) respectively, showed distinct patterns of host specificity at various developmental stages, and a higher fitness on their natural hosts, suggesting these races are locally adapted. Here we discuss the implications of our findings from a broader perspective. We suggest that differences in signal responsiveness and perception by the parasite, as well as qualitative differences in signal production by the host, may elicit host specificity in this parasitic plant. Together with our earlier demonstration that these O. minor races are genetically distinct based on molecular markers, our recent data provide a snapshot of speciation in action, driven by host specificity. Indeed, host specificity may be an underestimated catalyst for speciation in parasitic plants generally. We propose that identifying host specific races using physiological techniques will complement conventional molecular marker-based approaches to provide a framework for delineating evolutionary relationships among cryptic host-specific parasitic plants.Key words: host specificity, parasitic plant, broomrape, orobanche, speciation     

The gut bacterial community affects immunity but not metabolism in a specialist herbivorous butterfly

1. Plant tissues often lack essential nutritive elements and may contain a range of secondary toxic compounds. As nutritional imbalance in food intake may affect the performances of herbivores, the latter have evolved a variety of physiological mechanisms to cope with the challenges of digesting their plant‐based diet. Some of these strategies involve living in association with symbiotic microbes that promote the digestion and detoxification of plant compounds or supply their host with essential nutrients missing from the plant diet. In Lepidoptera, a growing body of evidence has, however, recently challenged the idea that herbivores are nutritionally dependent on their gut microbial community. It is suggested that many of the herbivorous Lepidopteran species may not host a resident microbial community, but rather a transient one, acquired from their environment and diet. Studies directly testing these hypotheses are however scarce and come from an even more limited number of species.
2. By coupling comparative metabarcoding, immune gene expression, and metabolomics analyses with experimental manipulation of the gut microbial community of prediapause larvae of the Glanville fritillary butterfly (Melitaea cinxia, L.), we tested whether the gut microbial community supports early larval growth and survival, or modulates metabolism or immunity during early stages of development.
3. We successfully altered this microbiota through antibiotic treatments and consecutively restored it through fecal transplants from conspecifics. Our study suggests that although the microbiota is involved in the up‐regulation of an antimicrobial peptide, it did not affect the life history traits or the metabolism of early instars larvae.
4. This study confirms the poor impact of the microbiota on diverse life history traits of yet another Lepidoptera species. However, it also suggests that potential eco‐evolutionary host‐symbiont strategies that take place in the gut of herbivorous butterfly hosts might have been disregarded, particularly how the microbiota may affect the host immune system homeostasis.

     

Plant–plant communication and community of herbivores on tall goldenrod

1. The volatiles from damaged plants induce defense in neighboring plants. The phenomenon is called plant–plant communication, plant talk, or plant eavesdropping. Plant–plant communication has been reported to be stronger between kin plants than genetically far plants in sagebrush.
2. Why do plants distinguish volatiles from kin or genetically far plants? We hypothesize that plants respond only to important conditions; the induced defense is not free of cost for the plant. To clarify the hypothesis, we conducted experiments and investigations using goldenrod of four different genotypes.
3. The arthropod community on tall goldenrods were different among four genotypes. The response to volatiles was stronger from genetically close plants to the emitter than from genetically distant plants from the emitter. The volatiles from each genotype of goldenrods were different; and they were categorized accordingly. Moreover, the arthropod community on each genotype of goldenrods were different.
4. Synthesis: Our results support the hypothesis: Goldenrods respond to volatiles from genetically close plants because they would have similar arthropod species. These results are important clues elucidating adaptive significance of plant–plant communication.

​     

Complete plastid genome sequences suggest strong selection for retention of photosynthetic genes in the parasitic plant genus <Emphasis Type="Italic">Cuscuta</Emphasis>

Background  

Plastid genome content and protein sequence are highly conserved across land plants and their closest algal relatives. Parasitic plants, which obtain some or all of their nutrition through an attachment to a host plant, are often a striking exception. Heterotrophy can lead to relaxed constraint on some plastid genes or even total gene loss. We sequenced plastid genomes of two species in the parasitic genus Cuscuta along with a non-parasitic relative, Ipomoea purpurea, to investigate changes in the plastid genome that may result from transition to the parasitic lifestyle.     

WARPP—web application for the research of parasitic plants

The lifestyle of parasitic plants is associated with peculiar morphological, genetic, and physiological adaptations that existing online plant-specific resources fail to adequately represent. Here, we introduce the Web Application for the Research of Parasitic Plants (WARPP) as an online resource dedicated to advancing research and development of parasitic plant biology. WARPP is a framework to facilitate international efforts by providing a central hub of curated evolutionary, ecological, and genetic data. The first version of WARPP provides a community hub for researchers to test this web application, for which curated data revolving around the economically important Broomrape family (Orobanchaceae) is readily accessible. The initial set of WARPP online tools includes a genome browser that centralizes genomic information for sequenced parasitic plant genomes, an orthogroup summary detailing the presence and absence of orthologous genes in parasites compared with nonparasitic plants, and an ancestral trait explorer showing the evolution of life-history preferences along phylogenies. WARPP represents a project under active development and relies on the scientific community to populate the web app’s database and further the development of new analysis tools. The first version of WARPP can be securely accessed at https://parasiticplants.app. The source code is licensed under GNU GPLv2 and is available at https://github.com/wickeLab/WARPP.

The WARPP online resource is a new, expandable, and interactive parasitic plant-specific data hub that provides online tools tailored to the peculiarities of parasitic angiosperms.     

Mechanisms of species‐sorting: effect of habitat occupancy on aphids' host plant selection

相似文献   

Three-dimensional reconstructions of haustoria in two parasitic plant species in the Orobanchaceae

Parasitic plants infect other plants by forming haustoria, specialized multicellular organs consisting of several cell types, each of which has unique morphological features and physiological roles associated with parasitism. Understanding the spatial organization of cell types is, therefore, of great importance in elucidating the functions of haustoria. Here, we report a three-dimensional (3-D) reconstruction of haustoria from two Orobanchaceae species, the obligate parasite Striga hermonthica infecting rice (Oryza sativa) and the facultative parasite Phtheirospermum japonicum infecting Arabidopsis (Arabidopsis thaliana). In addition, field-emission scanning electron microscopy observation revealed the presence of various cell types in haustoria. Our images reveal the spatial arrangements of multiple cell types inside haustoria and their interaction with host roots. The 3-D internal structures of haustoria highlight differences between the two parasites, particularly at the xylem connection site with the host. Our study provides cellular and structural insights into haustoria of S. hermonthica and P. japonicum and lays the foundation for understanding haustorium function.

Three-dimensional image reconstruction visualized the spatial organization of cell types in the haustoria of the Orobanchaceae parasitic plants Striga hermonthica and Phtheirospermum japonicum.     

Mitochondrial redox systems as central hubs in plant metabolism and signaling

Plant mitochondria are indispensable for plant metabolism and are tightly integrated into cellular homeostasis. This review provides an update on the latest research concerning the organization and operation of plant mitochondrial redox systems, and how they affect cellular metabolism and signaling, plant development, and stress responses. New insights into the organization and operation of mitochondrial energy systems such as the tricarboxylic acid cycle and mitochondrial electron transport chain (mtETC) are discussed. The mtETC produces reactive oxygen and nitrogen species, which can act as signals or lead to cellular damage, and are thus efficiently removed by mitochondrial antioxidant systems, including Mn-superoxide dismutase, ascorbate–glutathione cycle, and thioredoxin-dependent peroxidases. Plant mitochondria are tightly connected with photosynthesis, photorespiration, and cytosolic metabolism, thereby providing redox-balancing. Mitochondrial proteins are targets of extensive post-translational modifications, but their functional significance and how they are added or removed remains unclear. To operate in sync with the whole cell, mitochondria can communicate their functional status via mitochondrial retrograde signaling to change nuclear gene expression, and several recent breakthroughs here are discussed. At a whole organism level, plant mitochondria thus play crucial roles from the first minutes after seed imbibition, supporting meristem activity, growth, and fertility, until senescence of darkened and aged tissue. Finally, plant mitochondria are tightly integrated with cellular and organismal responses to environmental challenges such as drought, salinity, heat, and submergence, but also threats posed by pathogens. Both the major recent advances and outstanding questions are reviewed, which may help future research efforts on plant mitochondria.

Plant mitochondria are key components of redox homeostasis and play vital roles in regulating cellular metabolism, thereby affecting development and stress tolerance at the whole plant level.

Advances

• Improved quantitative MS-based approaches have accelerated the study of mitochondrial protein abundance, turnover and PTMs.
• Mitochondrial enzymes and cellular compartments operate interactively and efficiently exchange substrates.
• Roles for mitochondrial retrograde signaling in plant growth, during physiologically relevant stress conditions and in interaction with other organelles such as the chloroplasts, have been clarified.
• Further insights into mitochondrial antioxidant and peroxidase systems and how they affect other redox systems, enzymes, and whole plant growth have been generated.
• Our understanding of how mitochondria help plants power development and cope with adversity has improved.

     

Current progress in Striga management

The Striga, particularly S. he rmonthica, problem has become a major threat to food security, exacerbating hunger and poverty in many African countries. A number of Striga control strategies have been proposed and tested during the past decade, however, further research efforts are still needed to provide sustainable and effective solutions to the Striga problem. In this paper, we provide an update on the recent progress and the approaches used in Striga management, and highlight emerging opportunities for developing new technologies to control this enigmatic parasite.

Advances

• The recently established Striga control technologies, such as push-pull, toothpick, and imidazolinone seed dressing have opened up new opportunities for smallholder farmers to overcome this parasite.
• The development of low-cost and efficient germination stimulants together with an application protocol for rain-fed agriculture has made the suicidal germination strategy a realistic approach.
• Molecular elucidation of strigolactone biosynthesis and perception has led to the development of new chemicals that disrupt the communication between Striga and its hosts.

     

Plant phylogeny drives arboreal caterpillar assemblages across the Holarctic

1. Assemblages of insect herbivores are structured by plant traits such as nutrient content, secondary metabolites, physical traits, and phenology. Many of these traits are phylogenetically conserved, implying a decrease in trait similarity with increasing phylogenetic distance of the host plant taxa. Thus, a metric of phylogenetic distances and relationships can be considered a proxy for phylogenetically conserved plant traits and used to predict variation in herbivorous insect assemblages among co‐occurring plant species.
2. Using a Holarctic dataset of exposed‐feeding and shelter‐building caterpillars, we aimed at showing how phylogenetic relationships among host plants explain compositional changes and characteristics of herbivore assemblages.
3. Our plant–caterpillar network data derived from plot‐based samplings at three different continents included >28,000 individual caterpillar–plant interactions. We tested whether increasing phylogenetic distance of the host plants leads to a decrease in caterpillar assemblage overlap. We further investigated to what degree phylogenetic isolation of a host tree species within the local community explains abundance, density, richness, and mean specialization of its associated caterpillar assemblage.
4. The overlap of caterpillar assemblages decreased with increasing phylogenetic distance among the host tree species. Phylogenetic isolation of a host plant within the local plant community was correlated with lower richness and mean specialization of the associated caterpillar assemblages. Phylogenetic isolation had no effect on caterpillar abundance or density. The effects of plant phylogeny were consistent across exposed‐feeding and shelter‐building caterpillars.
5. Our study reveals that distance metrics obtained from host plant phylogeny are useful predictors to explain compositional turnover among hosts and host‐specific variations in richness and mean specialization of associated insect herbivore assemblages in temperate broadleaf forests. As phylogenetic information of plant communities is becoming increasingly available, further large‐scale studies are needed to investigate to what degree plant phylogeny structures herbivore assemblages in other biomes and ecosystems.

     

Root nematode infection enhances leaf defense against whitefly in tomato

The foliar response to different herbivores sharing the same hosts is an important topic for the study of plant-insect interactions. Plants evolve local and systemic resistant strategies to cope with herbivores. Many researchers have characterized the mechanisms of leaf responses to insect infestation; however, the fact that roots serve as systemic resistance modulators to leaf herbivores has been widely ignored. Here, we report that tomato (Solanum lycopersicum) plants infected with southern root-knot nematodes (Meloidogyne incognita)—which feed on the roots to form nodules—enhanced leaf defenses against aboveground attackers, specifically, the whitefly (Bemisia tabaci). Our results show that nematode infection reduced the whitefly population abundance because of conferring a stronger SA-dependent defense pathway against whitefly than in tomato plants without nematode infection. Meanwhile, nematode-infected tomato plant also activated the foliar JA-dependent defense pathway at 4 h after whitefly infestation. However, the foliar JA-dependent defense under whitefly infestation alone was suppressed, with the JA content being nearly 30 % lower than that in tomato plants co-infected with nematodes and whiteflies. Furthermore, nematode infection significantly decreased the plant nitrogen concentration in leaves and roots. As a result, nematode infection reduced the number of whiteflies by enhancing foliar SA-dependent defense, activating JA-dependent defense and decreasing nitrogen nutrition. Our results suggest that underground nematode infection significantly enhances the defense ability of tomato plants against whitefly.