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Schematic illustration of the structures of haustoria of Cuscuta campestris and Phtheirospermum japonicum. (A) Cuscuta campestris, a holoparasitic plant belonging to Convolvulaceae, develops lateral haustoria. (B) Phtheirospermum japonicum, a hemiparasitic plant belonging to Orobanchaceae, develops lateral haustoria. Holdfast of C. campestris and haustorial hair of P. japonicum are likely to be analogous that develop from epidermal cells and contribute to the adhesion of parasite to host. Intrusive cells of C. campestris, searching hyphae, develop from digitate cells which have been differentiated from the cortex or endodermal cells of the stem. On the other hand, intrusive cells of P. japonicum are shown to be differentiated from the epidermal cells.

Schematic illustration of the structures of haustoria of Cuscuta campestris and Phtheirospermum japonicum. (A) Cuscuta campestris, a holoparasitic plant belonging to Convolvulaceae, develops lateral haustoria. (B) Phtheirospermum japonicum, a hemiparasitic plant belonging to Orobanchaceae, develops lateral haustoria. Holdfast of C. campestris and haustorial hair of P. japonicum are likely to be analogous that develop from epidermal cells and contribute to the adhesion of parasite to host. Intrusive cells of C. campestris, searching hyphae, develop from digitate cells which have been differentiated from the cortex or endodermal cells of the stem. On the other hand, intrusive cells of P. japonicum are shown to be differentiated from the epidermal cells.

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Parasitic plants infect a broad range of plant species including economically important crops. They survive by absorbing water, minerals, and photosynthates from their hosts. To support their way of life, parasitic plants generally establish parasitic organs that allow them to attach to their hosts and to efficiently absorb substances from the vasc...

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... Understanding the molecular basis of parasitism by dodders is critical to develop effective pest control systems since they heavily damage many important crops. There are increasing number of studies about the development of the haustorium, a root-like structure that penetrates the host's tissue and draws water and nutrients (Yoshida et al. 2016;Shimizu and Aoki 2019; Jhu and Sinha 2022;Hartenstein et al. 2023). Elongation of the haustorium is promoted by (which was not certified by peer review) is the author/funder. ...
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The dodder, Cuscuta, is an obligate parasitic plant that cannot survive without a host plant and causes major damage to crop yields. To know its growth characteristics before parasitism, we examined the effect of various environmental conditions on seed germination and seedling growth in Cuscuta campestris. As for the effect of light on germination, far-red light was rather preferable to red light and the reversible response of the seeds to red and far-red light was confirmed, implicating a phytochrome-mediated signaling opposite to that in many seed plants. Among amino acids, aspartic acid and alanine had a promotive effect and histidine had an inhibitory effect on germination. We further found that, in addition to gibberellic acid, methyl jasmonate was stimulatory to germination and shoot elongation. While 2,4-D extended the viability of trichomes around the root cap, kinetin induced the formation of scale leaves on the shoot and calli at the base of the shoot and the root tip. RT-PCR experiments confirmed that expression of a putative RbcS gene for photosynthesis showed no response to light but that of a Phytochrome A homolog was increased in the dark. Our results indicate that some of the molecular mechanisms to respond to light and hormone signals are uniquely modified in dodder seedlings, providing a clue for understanding the survival strategy of parasitic plants.
... In Mexico, C. corymbosa has been reported from the state of Nayarit in the north to the state of Oaxaca in the south (McVaugh 1987). As a parasitic plant, C. corymbosa attaches itself to the host plants using specialized structures called haustoria, which penetrate the host's vascular tissue to extract water and nutrients (Shimizu and Aoki 2019). This parasitic relationship can weaken and ultimately damage the host plant, affecting its growth and productivity. ...
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The ever-growing consumption of herbs around the globe has motivated the researchers to acquire practical knowledge about other potential applications in human and animal health. In this research, an unmodified adsorbent prepared from the holoparasitic herb C. corymbosa was utilized for the removal of the carcinogen aflatoxin B1 (AFB1) from aqueous solutions. The adsorbent was characterized by Fourier transform near-infrared/mid-infrared spectrophotometry (FT-NIR/MIR), environmental scanning electron microscopy (ESEM), energy-dispersive X-ray fluorescence spectroscopy (EDX), X-ray diffraction (XRD), and point of zero charge (pHpzc). Adsorption experiments were carried out in batch systems, and the experimental data was used for isothermal (Langmuir and Freundlich) and kinetic (linear and non-linear forms of the pseudo-first and pseudo-second order) models. In general, the unmodified adsorbent removed AFB1 independent of the solution pH, showing a theoretical adsorption capacity of 555.76 mg AFB1/g at 303 K, significantly higher than that reported for other plant-based adsorbents and comparable with the efficiency of various inorganic adsorbents. Non-electrostatic attractions such as hydrogen bonding and dispersion forces along with complexation mechanisms were the primary interactions responsible for the adsorption of the pollutant. Our results clearly show that C. corymbosa could be a promising material for practical adsorption applications in the drinking water industry.
... Similar extracellular penetration of haustorium into host tissue has been reported by Dörr (1967), Nagar et al. (1984), Dawson et al, (1994), Vaughn (2003), Lee and Lee. (1991), Birschwilks et al. (2007), Lee (2009), andAoki (2019). A combination of physical pressure exerted by the elongating cells and enzymatic degradation are employed by the haustoria of Cuscuta to penetrate host tissues (Kuijt, 1969;Nagar et al. 1984;Dawson et al., 1994;Srivastava et al., 1994;Lee & Lee, 1989;López-Curto et al., 2006;Lee, 2007;Fathoulla & Duhoky 2008;Johnsen et al., 2015;Kaiser et al., 2015). ...
... This mechanism gives rise to a chimeric wall of the penetrating hyphae derived from both the host and the parasite. The intrusive cells have also been found to penetrate between the host cells by degrading the middle lamella and separating the cells (Vaughn, 2003;Lee & Lee, 1991;Lee, 2009;Shimizu & Aoki, 2019). These extracellular growth pathways are expected to minimize stress to the host (Vaughn, 2003). ...
... A few reports also mention intracellular growth of the unicellular searching hyphae within the host parenchyma (Birschwilks et al. 2007;Lee, 2009). Several investigations have revealed that parasitic plants directly secrete or induce secretion by hosts, substances that remodel cell walls of the host (Johnsen et al. 2015;Sarić-Krsmanović, 2019;Shimizu & Aoki, 2019) as well as parasite (Johnsen et al., 2015;Olsen & Krause, 2017) to enable penetration of the haustorium. ...
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Cuscuta reflexa Roxb. parasitises by developing and inserting a haustorium into host plant tissues. The development of haustorium in C. reflexa was studied in the host plants, Alstonia scholaris (L.) R.Br., Bougainvillea spectabilis Willd., Volkameria inermis L., and Senna siamea (Lam.) H.S. Irwin & Barneby. In all the hosts, haustorium development begins with the differentiation of a meristem from a few starch-containing cortical/pericycle cells that are adjacent to the parasite phloem tissue proximal to the host. The meristem cells divide and form files of parenchymatous cells which elongate and give rise to the haustorium. The distal-most file cells of the haustorium grow intrusively and penetrate through the parasite and host tissues. Thus, the haustorium is made up of elongated proximal files of ('axial') cells, and distal intrusive cells. The intrusive cells grow apically through the intercellular spaces and spaces created by the degeneration of cells in the path of the growing haustorium. Near the host vascular bundles, the intrusive cells of the haustorium diverge, grow towards, and make contact with the phloem and xylem tissues. The growth pattern of the endophytic haustorium was found to vary, based on the type of vasculature present in the four different hosts studied. In the stem of B. spectabilis, which possessed medullary, and a peripheral ring of vascular bundles, the haustorium grows deep into the pith between the bundles. The intrusive cells connect with numerous vascular bundles as the haustorium progresses into the pith. In the petiole of S. siamea and stems of V. inermis and A. scholaris, the vasculature consists of a ring of phloem tissue surrounding an inner ring of xylem tissue. Additionally, internal phloem is present inner to the primary xylem in A. scholaris. The haustorium in the first two hosts does not breach the cylinder of the xylem but in A. scholaris, the intrusive cells grow through the xylem ring and reach the internal phloem tissue. These findings indicate that the intrusive cells are strongly drawn to the host vascular tissues. Following contact with the host xylem, the intrusive cells differentiate into the xylem elements, and the differentiation continues into the axial cells of the haustorium. However, neither axial cells nor intrusive cells in contact with the host phloem were observed to differentiate into phloem cells or elements. In a mature haustorium, a continuous column of tracheary elements connecting the xylem of the host with that of the parasite is observed. Since only a few of the intrusive cells that are in contact with the host xylem tissue and only the central axial cells of the haustorium are found to differentiate into xylem elements, it is proposed that a two-way interaction between the signal/s derived from the host vascular cells and the cells of the haustorium, probably having procambium-like attribute, is a prerequisite for the differentiation of not only the intrusive cells but also the (procambium-like) cells of the haustorium into conducting cells, to establish a continuous vascular connection between the host and the parasite.
... Unlike root parasites, the obligate parasite dodders (Cuscuta) establish haustoria on the above-ground parts of their hosts [3,22]. In an early attempt to understand the genetic makeup of dodders, the expressed genes of the two Cuscuta species, C. pentagona and C. suaveolens, were analyzed by high-throughput sequencing [23]. ...
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Parasitic plants extract nutrients from the other plants to finish their life cycle and reproduce. The control of parasitic weeds is notoriously difficult due to their tight physical association and their close biological relationship to their hosts. Parasitic plants differ in their susceptible host ranges, and the host species differ in their susceptibility to parasitic plants. Current data show that adaptations of parasitic plants to various hosts are largely genetically determined. However, multiple cases of rapid adaptation in genetically homogenous parasitic weed populations to new hosts strongly suggest the involvement of epigenetic mechanisms. Recent progress in genome-wide analyses of gene expression and epigenetic features revealed many new molecular details of the parasitic plants’ interactions with their host plants. The experimental data obtained in the last several years show that multiple common features have independently evolved in different lines of the parasitic plants. In this review we discuss the most interesting new details in the interaction between parasitic and host plants.
... A pectin methylesterase enzyme was secreted from calli and seedlings of Orobranche aegyptiaca and O. cumana and it was speculated that the enzyme may have a function in host infection [51,117,118]. Additionally, a papain-like cysteine peptidase (cuscutain) has been identified in Cuscuta reflexa, involved in haustorium development and penetration [119,120]. The proteome revealed a metabolic enzyme related to glycoside hydrolases (pectinesterase) involved in complex polysaccharide degradation, and a DeSIlike protein similar to a peptidase, for which we performed measurements to obtain the different spatial activities of cellulases, xylanases, endo-1,4-β-glucanases, β-1,4-glucosidases, and peptidase-like proteins secreted by P. calyculatus during invasive stages on mesquite trees. ...
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Psittacanthus calyculatus is a hemiparasite mistletoe that represents an ecological problem due to the impacts caused to various tree species of ecological and commercial interest. Although the life cycle for the Psittacanthus genus is well established in the literature, the development stages and molecular mechanism implicated in P. calyculatus host infection are poorly understood. In this study, we used a manageable infestation of P. laevigata with P. calyculatus to clearly trace the infection, which allowed us to describe five phenological infective stages of mistletoe on host tree branches: mature seed (T1), holdfast formation (T2), haustorium activation (T3), haustorium penetration (T4), and haustorium connection (T5) with the host tree. Proteomic analyses revealed proteins with a different accumulation and cellular processes in infective stages. Activities of the cell wall-degrading enzymes cellulase and β-1,4-glucosidase were primarily active in haustorium development (T3), while xylanase, endo-glucanase, and peptidase were highly active in the haustorium penetration and xylem connection (T4). Patterns of auxins and cytokinin showed spatial concentrations in infective stages and moreover were involved in haustorium development. These results are the first evidence of proteins, cell wall-degrading enzymes, and phytohormones that are involved in early infection for the Psittacanthus genus, and thus represent a general infection mechanism for other mistletoe species. These results could help to understand the molecular dialogue in the establishment of P. calyculatus parasitism.
... These aspects have been reviewed recently (Kaiser et al., 2015;Kim and Westwood, 2015;Wu, 2018;. On the other hand, although review articles cover some specific aspects of Cuscuta haustorium development and parasitism behaviours (Yoshida et al., 2016;Shimizu and Aoki, 2019), several recent discoveries in the field have not been systematically reviewed yet. ...
... Haustorium organogenesis in Cuscuta species has been wellreviewed (Yoshida et al., 2016;Kokla and Melnyk, 2018;Shimizu and Aoki, 2019). However, the developmental stages are reported differently among references and diverse names are used to refer to the same stage, leading to some confusion. ...
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Parasitic plants are notorious for causing serious agricultural losses in many countries. Specialized intrusive organs, haustoria, confer on parasitic plants the ability to acquire water and nutrients from their host plants. Investigating the mechanism involved in haustorium development not only reveals the fascinating mystery of how autotrophic plants evolved parasitism but also provides the foundation for developing more effective methods to control the agricultural damage caused by parasitic plants. Cuscuta species, also known as dodders, are one of the most well-known and widely spread stem holoparasitic plants. Although progress has been made recently in understanding the evolution and development of haustoria in root parasitic plants, more and more studies indicate that the behaviors between root and stem haustorium formation are distinct, and the mechanisms involved in the formation of these organs remain largely unknown. Unlike most endoparasites and root holoparasitic plants, which have high host-specificity and self- or kin-recognition to avoid forming haustoria on themselves or closely related species, auto-parasitism and hyper-parasitism are commonly observed among Cuscuta species. In this review, we summarize the current understanding of haustorium development in dodders and the unique characteristics of their parasitizing behaviors. We also outline the advantages of using Cuscuta species as model organisms for haustorium development in stem holoparasitic plants, the current unknown mysteries and limitations in the Cuscuta system, and potential future research directions to overcome these challenges.
... Host localization is believed to involve both light [10] and chemical [11] signal perception. Formation of haustoria was recently reviewed by Shimizu and Aoki [12] and divided into three distinct phases. First, a tight adhesion is achieved through the secretion of adhesive substances and elongation of Cuscuta stem cells toward the host tissue. ...
... Various cell wall hydrolyzing and modifying enzymes are also involved in this process in order to loosen host cells' walls and facilitate the process [15,16]. Finally, during the conductive phase, the searching hyphae differentiate into xylem and establish a xylem (and probably phloem) bridge with the host [12]. ...
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Cuscuta australis is a widely distributed stem parasitic plant, infecting a variety of host plants. Its parasitism has a negative effect on the hosts, mainly due to the exhaustion of nutrients, thus negatively affecting the growth and development. However, recent studies indicated that the effect of parasitism may extend beyond the simple extraction of organic compounds, water, and minerals. In the present study, the model plant Arabidopsis thaliana was used as a host for Cuscuta australis, to study the effect of the parasite on the photosynthetic parameters and the proteome after short-term infection. To test this, a highly sensitive portable photosynthesis system and gel-based MS/MS proteomics were employed. It was found that the parasite has a dramatic negative effect on the photosynthetic ability of the host, as well as causing the up-regulation of stress-related proteins. Simultaneously, proteins involved in both decreased permeability and loosening of the cell wall of the host were found to be up-regulated.
... its leaves are reduced to vestigial scales (Mcneal et al., 2007). it is one of the most widespread and successful parasitic weeds parasitising a number of angiosperms (kujit, 1969;Malik & Singh, 1979;Dawson et al., 1994;Vaughn, 2002;Shimizu & aoki, 2019). Some Cuscuta species are "specialists" with a narrow host range while others like C. reflexa roxb. ...
... according to alakonya et al. (2012), the region of Cuscuta stem from which the meristem for haustorium development is derived has sufficient indeterminate meristematic activity to be able to form vegetative as well as flowering branches, and thus, may be involved in the regeneration of new shoots. the anticlockwise twining of C. reflexa stem around the host organs observed in the present study is also reported by tada et al. (1996), albert et al. (2008), Dey and Mukherjee (2013) and Shimizu and aoki, (2019), and appears to be a consistent feature. thick mats of yellow-green stands of C. reflexa are found generally on the terminal exposed branches of the trees with dense canopy such as Holoptelea integrifolia, Cascabela thevetia, Ziziphus mauritiana, Aegle marmelos, Alstonia scholaris covering large parts of host tree; very few strands are present on the lower branches. ...
... attachment of Cuscuta to host tissue is facilitated at the region of contact with the host, through the development of adhesive disks (Dörr, 1968;heide-Jørgensen, 1991;Dawson et al., 1994;Fathoulla & Duhoky, 2008), also called holdfasts (Shimizu & aoki, 2019) or pre/upper haustoria which include the haustorial meristem (kujit, 1977;lee & lee, 1989;Vaughn, 2002;lee, 2008;hong, 2011;alakonya et al., 2012;kaiser et al., 2015). although coiling of Cuscuta stem and haustorial initiation including prehaustoria formation can occur on plastic rods (Furuhashi et al., 2011), dead twigs and metal rods (kaiser et al., 2015), according to Fathoulla and Duhoky (2008), adhesive disks form only on contact with a live host. ...
Article
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Cuscuta reflexa roxb. (Convolvulaceae) is a total stem parasite that infects a large number of angiosperms. it grows and survives by parasitising host plants and extracting water and nutrients by developing and inserting into the host tissues, a haustorium. in general, C. reflexa stem infects the aerial parts of host plants. Field observations revealed that C. reflexa is found on the canopies of the plants well exposed to sunlight and often exhibits an annual cyclic pattern of growth on some trees such as Holoptelea integrifolia, Cascabela thevetia and Ziziphus mauritiana. it dries and withers in the months the hosts shed their leaves and regenerates after the hosts have sprouted new leaves. Moreover, C. reflexa is seen to forage and infect preferentially certain host plants such as H. integrifolia. The first step in parasitisation by C. reflexa is to attach tightly to the host plant by developing an adhesive disk at the site of contact with the host. the nutrient-sucking parasitic organ, the haustorium, forms in the center of the disk and penetrates the host tissues. Since the formation of adhesive disk is an important process in the parasitisation by Cuscuta, and there is a paucity of information on the subject, the process was investigated in a variety of host plants. the structural development of the adhesive disk of C. reflexa was studied in the widely related host plants, Alstonia scholaris (l.) r.Br., Bougainvillea spectabilis Willd., Volkameria inermis l. and Senna siamea (lam.) h.S. irwin & Barneby, using light microscopy. The process during parasitisation of the different host plants appears to be similar. at the site of contact with the host, the epidermal cells of the parasite along with a few underlying cortical cells enlarge and form the swollen adhesive disk. the epidermal cells elongate radially towards the host surface. they are vacuolated and possess hypertrophied nuclei. the outer apical surface of the cells in contact with host epidermis is highly invaginated/infolded and lobed. a layer of cementing material is found between the lobed epidermal surface of the disk and the outer surface of the host at the site of contact, binding the two surfaces tightly. the nature of the cementing material appears to be different from that of the primary cellulosic walls of the epidermis. The invaginated surface of the epidermal cells provides a larger interface for secretion of the cementing material enabling a tighter adhesion to the host.
... Dodder (Cuscuta spp.) is a vine plant with a parasitic stem without leaves and roots. It uses a unique organ, the haustorium (Yoshida et al., 2016), to attach to and penetrate the host's stem tissue forming a connection between the xylem and phloem and the host's vascular system (Shimizu and Aoki, 2019). The dodder parasitizes the plant host and steals nutrients and water from its vascular system, and exchanges a variety of molecules with it, including proteins, mRNAs, and small RNAs Shen et al., 2020;Song et al., 2022). ...
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Citrus is an essential horticultural fruit whose yield and quality are affected by salinity all over the world. The recognition and adaptive regulation of citrus against salt stress are important areas for cultivar improvement, but the vascular system signal transduction mechanism of the plant response to salt stress remains elusive. In this study, we constructed a dodder (Cuscuta spp.) linked Hamlin sweet orange (Citrus sinensis) plant community in which deliver a vascular signal through the dodder in response to salt stress. RNA-seq technology was used to analyze the gene expression profile of citrus leaves after salt treatment. The results showed that a vascular signal was transmitted to a dodder-linked host plant, triggering a transcriptional response to salt stress. However, the phenotypic and transudative ability of the dodder changed after 24 h. The salt treatment group (Group S) and the dodder-linked group (Group D) respectively contained 1,472 and 557 differentially expressed genes (DEGs). 454 of which were common to both groups. The results of our analysis revealed that the gene expression categories in Group D represented a highly consistent trend compared to the group S plants, indicating that the dodder-bridged vascular signals activated the stress-response of citrus leaves for transcriptomic reconfiguration. The KEGG pathway database and an analysis of key drivers revealed that phenylpropanoid biosynthesis, photosynthesis-antenna proteins, starch and sucrose metabolism, plant hormone signal transduction, circadian rhythm, and MAPK signaling pathways were significantly enriched as the critical genes during salt stress. A systemic signal in the dodder-bridged host significantly regulated abiotic stress-related secondary metabolic pathways, including those for phenylpropanoids, lignin, and lignans. The physiological indexes of photosynthetic intensity, respiration, and attractiveness among communities supported the transcriptional changes. Thus, our results indicate that salt stress-induced vascular system signals can be transmitted through the vascular system of a dodder linking citrus plants, revealing the genetic regulation and physiological changes of citrus leaves responding to plant stress signal transmission.
... Previous studies showed that the haustorial cells of dodder expanded, and the number of haustorial cells increased when the haustoria invaded host plants [25,26]. Auxin and cytokinin can induce haustoria formation under dark conditions [5,27]. ...
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Low R/FR irradiation can promote dodder haustorium formation on the host plant; however, the mechanisms underlying the process are still unknown. In this study, we compared the transcriptomic data during the formation of haustorium of Cuscuta chinensis on host plant Arabidopsisthaliana under low (R/FR = 0.1) versus high (R/FR = 0.2) R/FR irradiation at 12 h, 24 h and 72 h time points. The results show that low R/FR radiation significantly promoted the entanglement and haustorium formation. Transcriptome analysis showed that during the early stage of haustorium formation, low R/FR radiation significantly up-regulated ARR-A related genes and down-regulated peroxidase related genes compared with high R/FR radiation. Meanwhile, during the middle stage of haustorium formation, low R/FR treatment significantly increased the expression of genes related to pectinesterase (PE), polygalacturonase (PG) and pectin lyase (Pel) production, while, during the late stage of haustorium formation, peroxidase (Prx)-related genes were differentially expressed under different R/FR treatments. Overall, our findings show that a low R/FR ratio promotes the parasitism of C. chinensis through plant hormone signal transduction and cell wall degradation pathways. This study provides a basis for the control of parasitic plants.