Trade winds are a widespread horticultural consideration throughout the tropics. Growth and productivity of most horticultural crops are not optimal on sites that are exposed to these chronic, unidirectional winds. We conducted four container studies on an exposed site, using clear plastic or screening material to provide three levels of wind exposure: 0%, 36%, or 100%. Two studies were conducted with direct-seeding, such that seedling emergence and early growth were determined for 7 weeks. Two studies were conducted using 8-week-old nursery plants that had been grown in a protected nursery. These plants were transplanted to the experimental site and grown for 6 weeks. Cultivars were `Known You 1', `Sunrise', and `Tainung 2'. Full exposure to wind reduced height up to 32%, increased root: canopy ratio up to 36% and exhibited no influence or slightly reduced stem cross-sectional area when compared with full protection from wind. Net carbon dioxide assimilation (Pn) was measured on intervals of about 2 h throughout several 24-h periods. Although the daily pattern depended on cultivar and date, the general trend was for Pn to be unaffected by wind from early to mid-morning, and for Pn of the unprotected plants to decline below that of the protected plants throughout the rest of the day. The Pn of plants receiving intermediate protection was highly variable among the cultivars and dates in relation to the protected and unprotected plants. Moreover, dark respiration of the unprotected plants was greater than the protected plants throughout the entire nocturnal period. The primary influence of wind on growth of young papaya seedlings was a shift in biomass allocation in favor of the stem base and roots.
Hiphil S. Clemente and Thomas E. Marler
Thomas E. Marler and Mark A. Lander
The frequency of tropical cyclones is a major factor affecting the vegetation of the Mariana Islands, where these storms are called typhoons. An average of about one typhoon per year has passed within ≈100 km of Guam during the past 50 years. The physiognomy of Guam's natural and urban forests is largely determined by these typhoons. The impact of each typhoon is determined by a long list of interacting factors such as species characteristics; environmental and horticultural conditions preceding the typhoon; the intensity, direction, and duration of winds; the amount of rainfall associated with the typhoon; and the environmental and horticultural conditions following the disturbance. Many species survive typhoons by reducing aerodynamic drag of the canopy by abscising inexpensive leaves or breakage of small stems which results in an intact major structural framework. Speed of recovery for nonlethal damage following disturbance depends on nonlimiting conditions during recovery. Thus, the most destructive typhoons are those that occur in sequence with other environmental stresses. The most common of these may be heat and high-light stress, associated with subsequent high pressure systems, and severe drought conditions. For example, the 230–298 km·h–1 winds of Typhoon Paka in Dec. 1997 were followed by the driest year on record for Guam. Typhoon debris and drought generated 1400 forest and grassland fires from Jan. through May 1998. Sequential typhoons are also severely damaging. For example, Guam experienced three direct eye passages and two more typhoons within 113 km during the months Aug. to Nov. 1992. Damage susceptibility and recovery dynamics will be discussed in relation to these and other physical, chemical, biological, and human-induced factors.
Thomas E. Marler and Michael V. Mickelbart
The influence of drought stress on leaf gas exchange and chlorophyll fluorescence characteristics of field-grown papaya (Carica papaya L.) plants was determined under a range of incident light fluxes and times of day. These data may aid in improving management systems for papaya production which minimize detrimental effects from suboptimal environmental conditions. Water was withheld from field-grown `Red Lady' plants in one study and `Tainung #2', `Red Lady', and `Sunrise' plants in a second study until soil matric potential was -60 to -70 kPa. Drought-stressed plants exhibited reduced net CO2 assimilation (ACO2) above light saturation, photosynthetic photon flux (PPF) at which light saturation for ACO2 occurred, and apparent quantum yield compared to well-watered plants. The light compensation point of drought-stressed plants was greater than that of well-watered plants. Leaf chlorophyll fluorescence characteristics were not influenced by drought stress. The daily pattern of leaf gas exchange was dependent on climatic conditions. For sunny days, ACO2, stomatal conductance of water (gs), and water use efficiency of well-watered plants were maximal at mid-morning, declined during midday, and then partially recovered during late afternoon. In drought-stressed plants, leaf gas exchange was relatively constant after a brief early morning maximum. On overcast days, the responses of gas exchange variables in relation to time of day followed smooth bell-shaped patterns regardless of the level of drought stress. Combined with previously published data, these results indicate that the influence of drought stress on gas exchange is highly dependent on time of day, ambient sky conditions, plant size, and speed with which drought stress occurs.
Thomas E. Marler and Frederick S. Davies
Growth responses of young `Hamlin' orange [Citrus sinensis (L.) Osbeck] on sour orange (C. aurantium L.) trees to microsprinkler irrigation were studied under field conditions from 1985 to 1987 to determine the most-efficient irrigation rates and duration. Trees were irrigated when available soil water depletion (SWD) reached 20% (high frequency), 45% (moderate frequency), and 65% (low frequency). Trees at the moderate and low levels received 49% and 13%, respectively, as much irrigation water as the high treatment. Canopy volume, trunk cross-sectional area, dry weight, shoot length, leaf area, total root dry weight and volume, and new root dry weight were similar for the high and moderate levels in 2 of 3 years, but were significantly reduced at the low level. Summer and fall growth flushes were delayed or did not occur at the moderate and low levels. More than 90% of root dry weight was within 80 cm of the trunk at the end of the first growing season.
Thomas E. Marler and Patrick D. Lawton
Leaflets of `Arkin', `B-10', `Kary', and `Sri Kembangan' carambola (Averrhoa carambola L.) trees were restrained in a horizontal position for 3.5 h during midday under full sun conditions to determine the influence of overriding natural leaflet movement on adaxial chlorophyll fluorescence and temperature. Induced chlorophyll fluorescence obtained after 30 minutes of dark adaptation following the period of full sun exposure was affected by leaflet movement. Restrained leaflets exhibited a variable fluorescence (Fv)/peak fluorescence (Fm) of 0.48, while that of unrestrained leaflets was 0.65. Adaxial leaflet temperature of restrained leaflets was 6C higher than that of leaflets that were allowed to move. The influence of leaflet movement on temperature or chlorophyll fluorescence was not different among the four cultivars. However, mean Fv/Fm of `Kary' and `Sri Kembangan' was lower than that of `B-10'. Our results indicate that the ability of carambola to change leaflet angle leads to lower temperature and higher photochemical efficiency than occurs when leaflets are not allowed to move naturally (vertically orient) under full sun conditions.
Hiphil S. Clemente and Thomas E. Marler
Field-grown `Red Lady' papaya (Carica papaya L.) plants were used to measure foliar gas-exchange responses to rapid changes in irradiance levels to determine if papaya stomata are able to track simulated sun-to-cloud cover transitions. Natural sunlight and neutral shade cloth placed over the leaf were used to provide high photosynthetic photon flux (PPF) of about 2000 μmol·m-2·s-1 until leaves reached steady state within the cuvette, followed by three minutes with low PPF of about 325 μmol·m-2·s-1, and a return to PPF of about 2000 μmol·m-2·s-1. Net CO2 assimilation (A) declined from an initial 20 μmol·m-2·s-1 to about 9 μmol·m-2·s-1 within 20 seconds of initiating low PPF, and remained fairly stable for the duration of the three minutes of low PPF. Stomatal conductance (gs) declined within 60 seconds of initiating low PPF, from 385 to about 340 μmol·m-2·s-1 during the three minutes duration of low PPF. Following the return to high PPF, A rapidly increased to about 18 μmol·m-2·s-1, then gradually increased to the original value. After a lag of about 1 minute following the return to high PPF, gs began to increase and returned to the original value after three minutes. Container-grown `Tainung #1' papaya plants were used in a second study to determine the influence of mild drought stress on gas-exchange responses to rapid irradiance transitions. For drought-stressed plants, gs declined to a greater magnitude following the high-to-low PPF transition, and gs and A recovered more slowly following the transition from low-to-high PPF than for well-watered plants. Water use efficiency declined to a minimum immediately following the high-to-low PPF transition for both sets of plants, but recovered more rapidly for drought-stressed plants. These results indicate that papaya stomata are able to track rapid changes in irradiance, and mild drought stress enhances the tracking response.
Thomas E. Marler and Haluk M. Discekici
A laboratory exercise is outlined in which breath is used as the source for elevating CO2. Single-plant enclosures are constructed by placing containers with stem cuttings within clear bags for maintaining high humidity during root initiation. These enclosures provide a restricted atmosphere in which elevated CO2 is adequately confined. The materials are inexpensive and readily available. The procedure is rapid, with results obtained in as few as 7 days. The increase in canopy and/or root growth as a result of elevated CO2 may be observed visually by younger audiences, or may be measured and analyzed in a manner dependent on the age group. The exercise generates an abundance of discussion and may be used to lead into many lectures on plant physiology, horticultural crop production, or global concepts of CO2 and the plant kingdom.
Thomas E. Marler and Gil N. Cruz
Ubiquitous Aulacaspsis yasumatsui Takagi infestations of Guam’s Cycas micronesica K.D. Hill trees cause direct herbivory of most exposed organ surfaces, including developing naked ovules and seeds. The nonstructural carbohydrates of infested vs. noninfested seeds were quantified to understand more fully the influences on seed quality for propagation purposes. Two studies compared seeds from healthy trees with those of unhealthy trees suffering from whole-tree A. yasumatsui infestations. The sugars fructose, glucose, maltose, and sucrose were in greater concentrations and herbivory reduced these free sugars by a greater percentage in sarcotesta tissue than in sclerotesta and gametophyte tissues. Starch concentration was greatest in gametophyte tissue, but herbivory reduced starch by a greater percentage in sarcotesta tissue. A third study was used to manipulate seeds of unhealthy infested trees such that some seeds were uninfested and some seeds were infested and revealed the nonstructural carbohydrates of the uninfested seeds were greater than those of the infested seeds in patterns that were similar to those when the entire tree was protected from herbivory. The combined results indicated that both source and sink relations were involved in the reductions of seed carbohydrates by A. yasumatsui herbivory. The reduction in seed resource pool by the herbivore feeding may be one of the mechanisms that results in reduced germination percentage and increased seedling mortality.
Hiphil S. Clemente and Thomas E. Marler
Container-grown papaya plants were subjected to a slow drying cycle by replenishing a portion of the mean daily water loss. When light-saturated net CO2 assimilation (A) of stressed plants was ≈70% of well-watered plants, sun–cloud transitions were simulated by placing shadecloth between a lamp and the leaf cuvette. The cuvette was initially positioned to receive photosynthetic photon flux (PPF) of ≈1600 μmol–m–2–s–1. The shadecloth was placed over the cuvette for a 3-min period (250 μmol–m–2–s–1), then removed. Gas-exchange responses were recorded for another 3 min of high light. Within 20 sec of initiating low light on well-watered plants, A was <50% of the high-light value. Stomatal conductance (gs) of these plants began to decline by 60 sec and slowly declined to <90% of the high-light value. Both A and gs recovered to the original values by ≈2 min following return to the high-light conditions. For stressed plants, the decline of gs under low light began earlier and was of greater magnitude, and gs and A recovery following the return to high light was delayed. These results indicate that papaya has suntracking capabilities, and mild drought stress enhances the tracking response.
Thomas E. Marler and Christopher A. Shaw
The relationship between mature Cycas micronesica K.D. Hill seed sterol concentration and content and plant or seed phenotypic characteristics was established by multiple regression. Combined models were significant for free but not glycosylated sterols. Reduced models revealed leaf number as the only significant predictor. Free and glycosylated sterol concentrations were unaffected throughout the range of several predictors: tree height (1.7 to 5.8 m), seed fresh weight (48 to 120 g), seed load (one to 76 seeds per plant), and estimated tree age (32 to 110 years). The free and glycosylated sterol phenotypes were also not dependent on the presence/absence of developed embryos in mature seeds. The significant response to leaf number was subtle with an increase of 43 leaves associated with a 0.1-mg increase in free sterol per gram seed fresh weight. This is the first report for any cycad that discusses reproductive or physiological traits in the context of allometric relations. Results indicate a highly constrained phenotypic plasticity of Cycas gametophyte sterol and steryl glucoside concentration and seed content in relation to whole plant and organ size variation.