THE LEAF OF CALYCADENIA AND ITS GLANDULAR APPENDAGES

Carlquist , Sherwin . (Rancho Santa Ana Botanic Garden, Claremont, Calif.) The leaf of Calycadenia and its glandular appendages. Amer. Jour. Bot. 46(2) : 70-80. Illus. 1959.—Large tack-shaped glands are characteristic of the leaves of Calycadenia which are associated with the inflorescence. These glands may be divided into those which are terminal on leaves and those which occur laterally on the surface of the leaf. Lateral glands show stages early in their development which are identical with those of simpler trichomes of Madinae. Terminal glands, which possess more vascularization of the stalk, show a more modified form of development. Vascularization is not derived from protoderm, but from more deeply-seated cells. These cells are included in a zone of elongation which forms the stalk. Vascular bundles may extend to the base of glands which lack vascularization in their stalks. Tack-shaped glands are considered an advanced form of trichome in which internal tissues of the leaf are involved. Within the genus Calycadenia, ontogenetic and comparative studies suggest that the following characters are advanced: reduction to a single terminal gland, “inrolling” of margins to form a cylinder of bundles, concomitant with a central core of fibers or a pectic channel. Systematic distribution of gland occurrence and of types of foliar structure are given.     

ONTOGENY AND COMPARATIVE ANATOMY OF THORNS OF HAWAIIAN LOBELIACEAE

Carlquist , Sherwin . (Claremont Graduate School, Claremont, Calif.) Ontogeny and comparative anatomy of thorns of Hawaiian Lobeliaceae. Amer. Jour. Bot. 49(4): 413–419. Illus. 1962.—Species of Rollandia and Cyanea (sections Genuinae and Palmaeformes), endemic Hawaiian genera of Lobeliaceae, are unique in the family in possessing thorns and thorn-like structures on leaves, and in some cases, on stems and flowers. These thorns always originate in conjunction with a unicellular, non-glandular trichome which terminates the thorn. Ontogenetic studies show that divisions leading to the formation of the thorn occur in the ground meristem as soon as the trichome is differentiated. Periclinal divisions predominate at first, but anticlinal and diagonal ones are also present at all stages. Thick secondary walls are formed on the trichome and other epidermal cells near the thorn tip. Periderm forms on old thorns of stems. Vascular tissue and laticifers are absent in thorns. Thorns in Cyanea and Rollandia seem best interpreted as specializations within these genera.     

Hair,encoding a single C2H2 zinc‐finger protein,regulates multicellular trichome formation in tomato

Trichomes originate from the epidermal cells of nearly all terrestrial plants, which are specialized unicellular or multicellular structures. Although the molecular mechanism regulating unicellular trichome formation has been extensively characterized, most of the genes essential for multicellular trichome formation remain unknown. In this study, we identified an associated locus on the long arm of chromosome 10 using a genome‐wide association study (GWAS) on type‐I trichomes of 180 diverse Solanum lycopersicum (tomato) accessions. Using map‐based cloning we then cloned the key gene controlling the initiation of this type of trichome, named Hair (H), which encodes a single C2H2 zinc‐finger protein. Transgenic experiments showed that hair‐absent phenotype is caused by the deletion of the entire coding region of H. We identified three alleles of H containing several missense mutations and a nucleotide deletion, which result in amino acid substitutions and a reading frame shift, respectively. In addition, knockdown of H or Woolly (Wo) represses the formation of type‐I trichomes, suggesting that both regulators may function as a heterodimer. Direct protein–protein interaction between them was further detected through pull‐down and yeast two‐hybrid assays. In addition, ectopic expression of H in Nicotiana tabacum (tobacco) and expression of its homologs from Capsicum annuum (pepper) and tobacco in tomato can trigger trichome formation. Taken together, these findings suggest that the H gene may be functionally conserved in multicellular trichome formation in Solanaceae species.     

Solitary terminal cells of Aphanizomenon gracile (Cyanobacteria,Nostocales) can divide and renew trichomes

An attempt was made to find evidence that morphologically distinct terminal cells of filamentous cyanobacterium Aphanizomenon gracile strain CCALA 8 are capable of dividing and forming trichomes. Based on our current knowledge, the division of morphologically diversified terminal cells is possible in nostocalean cyanobacteria. However, this process has been observed only in a few species. Terminal cells of A. gracile differ morphologically from other vegetative cells of a trichome, as they are not hyaline and can sometimes be found as solitary cells in cultures. Hence, it was reasonable for us to suspect that these cells are capable of dividing and forming trichomes. We observed terminal cells under a light and transmission electron microscope. Microscopic observations revealed that the septum formed in both solitary terminal cells and in terminal cells attached to trichomes. Our study is the first to demonstrate division and renewal of trichomes in terminal cells of A. gracile. Previously, such mode of reproduction was described only for another nostocalean cyanobacterium Raphidiopsis mediterranea. Moreover, our findings further emphasize the variability among members that belong to the genus Aphanizomenon , in which a type species (A. flos‐aquae) has hyaline cells incapable of dividing and renewing trichomes, while A. gracile can additionally propagate by solitary terminal cells division. This additional feature distinguishing A. gracile from typical species of Aphanizomenon, such as A. flos‐aquae, might be valuable for resolving taxonomic position of the species considering ambiguous genetic relationship between A. gracile and A. flos‐aquae.     

MORPHOLOGICAL AND 16S rRNA GENE EVIDENCE FOR RECLASSIFICATION OF THE PARALYTIC SHELLFISH TOXIN PRODUCING APHANIZOMENON FLOS‐AQUAE LMECYA 31 AS APHANIZOMENON ISSATSCHENKOI (CYANOPHYCEAE)1

A taxonomic reevaluation of the paralytic shellfish toxin (saxitoxins) producing cyanobacterium Aphanizomenon flos‐aquae Ralfs ex Born. & Flah. LMECYA31 was done using morphology and 16S rRNA gene sequences. We found that strain LMECYA31 was incorrectly identified as Aph. flos‐aquae based on (a) lack of bundle formation in trichomes, (b) shape of terminal cells in the trichomes, (c) lower similarity (<97.5%) in the 16S rRNA gene sequences relative to those of Aph. flos‐aquae, and (d) comparison within a phylogenetic tree of 16S rRNA gene sequences. The shape of the terminal trichome cells and the shape and size of the vegetative cell, heterocyst, and akinete in strain LMECYA31 match characters of Aph. issatschenkoi (Ussachew) Proschkina‐Larvernko. 16S rRNA gene sequences and phylogenetic clusters constructed from 16S rRNA gene sequences support our conclusion that strain LMECYA31 should be Aph. issatschenkoi.     

EVALUATION OF RELATIONSHIPS WITHIN ERIODICTYON (HYDROPHYLLACEAE) USING TRICHOME CHARACTERISTICS

All 13 taxa of Eriodictyon Bentham (Hydrophyllaceae) were examined to determine the range of variation in trichome characters within the genus. Four simple trichome types were found: short and straight, intermediate length and straight, long and straight, long and wavy. Glandular capitate trichomes were also found in some species. Sessile glands were also observed but not included in the study. Most taxa displayed unique combinations of trichome types on stems, leaves, inflorescence axes and flower parts that allowed those taxa to be identified using trichome types alone. Trichome data support most previous taxonomic treatments, but suggest that 1) the two varieties of E. traskiae are extremely similar, 2) the rare E. altissimum is most similar to the widespread E. californicum, 3) the rare E. capitatum is allied with E. crassifolium var. nigrescens or perhaps E. angustifolium, 4) E. crassifolium var. denudatum and E. crassifolium var. nigrescens are essentially indistinguishable using trichome characters and, coupled with variation in other characters, are best combined under the name E. crassifolium var. nigrescens. Trichome characters provide a wealth of taxonomically useful information and may prove useful in the study of related genera of Hydrophyllaceae.     

Plant cells which aid in pollen digestion within a beetle's gut

Summary The peach palm, Bactris gasipaes H.B.K., in Costa Rica, possesses unusual trichomes on the inflorescence epidermal surface. Certain cells of the trichome possess a thick, highly lignified cell wall and are consumed by the beetle Cyclocephala amazona L. before it ingests pollen from the same inflorescence. Chemical analyses show the trichome to possess no nutritive value. The thick-walled trichome cells pass intact through the beetle's digestive system, while ingested pollen is crushed. We suggest that the specialized plant cells function as gastroliths in the beetle's digestive tract.     

Geometry and mechanics of the morphogenesis of active membranes on the example of plant trichome cells of the genus <Emphasis Type="Italic">Draba</Emphasis> L.

The stages of the early morphogenesis of simple (unbranched) and complex (branched) unicellular trichomes are studied in two species of the genus Draba—D. sibirica (Pall.) Thell. and D. daurica DC. The geometry of morphogenesis is estimated by analyzing intraindividual variation of quantitative morphological characteristics of the developing leaf blade and peduncle trichomes. The surface of all types of trichome cells first acquires a spherical shape, followed by a U-shaped configuration with cylindrical proximal and spherical distal regions. In the development of complex trichomes, the area of the distal zone grows at a higher rate, which leads to separation of its volume into individual spherical regions, the morphogenesis of which repeats the early morphogenetic stages of the overall trichome cell, forming simple (unbranched) or complex (branched) trichome rays. As a rule, the lateral polarity of a trichome cell coincides with the proximodistal polarity of the leaf. Quantitative morphological data make it possible to infer an algorithm of the changes in shape common for all trichome cells, namely, the growth cycle comprising alternation of the phases of increase and decrease in the curvature of the outer cell surface. This surface is an active membrane expanded by the internal pressure and concurrently capable of actively increasing its area by incorporation of new structural elements. A distinctive feature of the proposed model is the geometrical inhomogeneity of the surface movement, changing the radius of curvature and creating internal (active) mechanical stresses in this membrane. A decrease in the ratio of the membrane surface area to the volume deprives the spatially homogeneous shape of its stability; correspondingly, the transition from elastic resistance to internal pressure to active resistance with the help of curvature differentiation becomes more energetically favorable. The source for growth and morphogenesis of the active membrane is alternation of the phases of local curvature leveling, which “charges” the membrane with active mechanical stresses and “discharge” of these stresses, leading to differentiation of the membrane’s local curvatures.     

Tissue organisation,pollen receptivity and pollen tube guidance in normal and mutant stigmas of the grass Pennisetum typhoides (Burm.) Stapf et Hubb.

Summary The normal stigma of Pennisetum typhoides is twin-branched, each branch bearing unbranched trichomes. As is general among the grasses, the papillate cells of the trichomes possess a discontinuous cuticle with overlying protein and polysaccharide secretions. These adaptations for pollen capture and hydration are absent from the stigma axes. Pollen tubes emerging from grains received on the trichomes are guided into the axes with the tips directed towards the ovary by the architecture of the basal cell complex. There are no defined transmitting tracts in the stigma axes, and further passage is through intercellular spaces of a tissue of elongated cells between the epidermis and the central vascular strands. In the mutant tr, tr, the stigmas are twin-branched, but lack trichomes. However, the principal adaptations of the trichome cells for the capture and hydration of pollen are expressed in the epidermal cells of the branches, which have permeable cuticles and the characteristic surface secretions. Pollen tubes emerging from grains germinating on the branches enter between the files of epidermal cells, or at their ends. In the absence of the guidance provided by trichome structure in the normal stigma, they pass indifferently either towards or away from the ovary. The implications of the comparison between the normal and mutant genotypes for understanding the requirements for pollen capture, germination and pollen-tube guidance in the grasses are discussed.     

TOMATO TRICHOMES AND MUTATIONS AFFECTING THEIR DEVELOPMENT

A new trichome type for the genus Lycopersicon is described in L. esculentum Mill. It is a short (0.03–0.08 mm), pendant, glandular hair with a club-shaped head consisting of 8–12 cells. Two previously described “hairless” mutations were examined microscopically. One, hl, does not affect the frequency of hairs nor the number of cells per hair, but causes abnormal enlargement of the stalk cells of all hair types, and thus produces shortened, extremely bent and twisted hairs. Observations on the time of action of this gene indicate that in trichome development two to four cell divisions occur prior to any appreciable cell enlargement. The second mutation, h, affects only the large type of trichome. This mutation effects a developmental shift from trichome to stomatal apparatus at the apex of the multicellular base normally supporting the large trichomes.     

Leaf epidermal features of Lithocarpus (Fagaceae) from China and their systematic significance

The leaves of 52 species of Lithocarpus in China were studied. The adaxial leaf epidermis was investigated by light microscopy. Epidermal cells of the adaxial surface were classified into three types on the basis of the outline of their anticlinal walls, i.e. sinuate, straight and curved. The abaxial leaf epidermis was investigated by light microscopy and scanning electron microscopy. The following types of trichome were observed: appressed parallel tuft, stellate, fused stellate, papillae, stipitate fasciculate, solitary unicellular, appressed laterally attached unicellular, curly thin‐walled unicellular, bulbous and thin‐walled peltate. The fused stellate, appressed laterally attached unicellular and curly thin‐walled unicellular trichomes were reported in Lithocarpus for the first time. The appressed parallel tuft trichome, which is recognized as a salient characteristic of Lithocarpus, was not found in 15 species. A cladistic analysis was performed on the basis of the leaf epidermal features. According to the leaf epidermal features and several morphological characteristics, 26 of the 52 species could be divided into seven groups. Similar groups can be found in Barnett's and Camus' systems. The trichomes of four genera in Fagaceae are listed and compared. Lithocarpus had 14 types of trichome, 11 of which were identical to types found in Quercus, more than in Castanopsis and Cyclobalanopsis. The evolutionary trends of trichomes in Fagaceae are discussed and a new point of view is raised. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 216–228.     

Trichome production and variation in young plant resistance to the specialist insect herbivore Plutella xylostella among natural populations of Arabidopsis lyrata

The strength of plant‐herbivore interactions varies spatially and through plant ontogeny, which may result in variable selection on plant defense, both among populations and life‐history stages. To test whether populations have diverged in herbivore resistance at an early plant stage, we quantified oviposition preference and larval feeding by Plutella xylostella (L.) (Lepidoptera: Plutellidae) on young (5–6 weeks old) Arabidopsis lyrata (L.) O'Kane & Al‐Shehbaz (Brassicaceae) plants, originating from 12 natural populations, six from Sweden and six from Norway. Arabidopsis lyrata can be trichome‐producing or glabrous, with glabrous plants usually receiving more damage from insect herbivores in natural populations. We used the six populations polymorphic for trichome production to test whether resistance against P. xylostella differs between the glabrous and the trichome‐producing morph among young plants. There was considerable variation among populations in the number of eggs received and the proportion of leaf area consumed by P. xylostella, but not between regions (Sweden vs. Norway) or trichome morphs. Rosette size explained a significant portion of the variation in oviposition and larval feeding. The results demonstrate that among‐population variation in resistance to insect herbivory can be detected among very young individuals of the perennial herb A. lyrata. They further suggest that trichome densities are too low at this plant developmental stage to contribute to resistance, and that the observed among‐population variation in resistance is related to differences in other plant traits.